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RIBOSOMAL AND CHLOROPLAST DNA EVIDENCE FORDIVERSIFICATION OF WESTERN AMERICAN PORTULACACEAE IN THE

ANDEAN REGION

EVIDENCIA DE ADN RIBOSOMAL Y DEL CLOROPLASTO PARA LADIVERSIFICACION DE LAS PORTULACACEAS DE AMERICA OCCIDENTAL

EN LA REGION ANDINA

Mark A. Hershkovitz1

1Departamento de Ciencias Ecológicas, Facultad de Ciencias, Universidad de Chile, Las Palmeras 3425, Santiago,Chile. [email protected].

ABSTRACT

Sequences of the ribosomal DNA internal transcribed spacer (ITS) region and the chloroplast DNA ycf3-trnS intergenicspacer were determined for 183 samples representing Chilean and non-Chilean taxa of western American Portulacaceaeand their outgroups. The data refine previous inferences of generic circumscriptions and interrelations. In particular, thedata reveal that an earlier circumscription of Cistanthe Spach is polyphyletic and also reveal a North American cladecomprising Claytonia L., Lewisia Pursh, Lewisiopsis R. Govaerts, and Montia L. Within the South American genera, twopatterns emerge from the data: (1) in some cases, interspecific divergence is remarkably low given the markers employedand estimated number of species; and (2) where divergence of one or both markers offer phylogenetic resolution, there isconflict between them and/or with morphology. The patterns can be evaluated in terms of two phenomena: (1)morphological/ecological radiation proceeding much faster than sequence divergence; and (2) frequent hybridization. Inthe latter case, the gene tree patterns may distort the true timing and cladistic pattern of morphological and ecologicaldiversification. At present, the degree to which evidence for hybridization among the Chilean Portulacaceae will prove tobe the rule or the exception is unclear. Nonetheless, spatial and temporal ecological patterns in Chile generally favorhybrid formation and persistence.

KEYWORDS: Portulacaceae, Chile, ITS, ycf3-trnS, hybridization.

RESUMEN

Se determinaron las secuencias del espaciador interno (ITS) del ADN ribosomal y del espaciador intergénico de ycf3-trnSdel ADN del cloroplasto para 183 muestras de taxas, chilenos y no chilenos, de Portulacaceae de América occidental, y susgrupos externos. Los datos permiten refinar previas inferencias sobre la circunscripción e interrelaciones de los génerosestudiados. En particular estos datos revelan que una circunscripción anteriormente publicada de Cistanthe Spach espolifilética y también revelan la existencia de un clado norteamericano compuesto por Claytonia L., Lewisia Pursh,Lewisiopsis R. Govaerts, and Montia L. De los datos de los géneros sudamericanos surgieron dos patrones: (1) en algunoscasos la divergencia interespecífica es notablemente baja dado los marcadores empleados y el número estimado deespecies; y (2) en cuanto la divergencia de uno o de los dos marcadores ofrecen resolución filogenética, existen conflictosentre ellos y/o entre la morfología. Estos patrones pueden ser interpretados en términos de dos fenómenos: diversificaciónmorfológica/ecológica siguiendo más rápida que la divergencia en las secuencias; y (2) frecuente hibridación. Este altogrado de hibridación sugiere que el árbol filogenético molecular puede representar mal el patrón temporal, morfológico yecológico de diversificación de estas especies. Al presente no es claro si la hibridación es una regla o una excepción entrelas Portulacaceae chilenas. No obstante, en Chile las condiciones ecológicas, espaciales y temporales, son en generalfavorables para la formación y persistencia de híbridos.

PALABRAS CLAVES: Portulacaceae, Chile, ITS, ycf3-trnS, hibridización.

Gayana Bot. 63(1): 13-74, 2006 ISSN 0016-5301

Gayana Bot. 63(1), 2006

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INTRODUCTION

The term “western American Portulacaceae” refers toa group of ca. 120-150 species of closely interrelatedgenera of traditional Portulacaceae (e.g., sensu Carolin1993) distributed predominantly in the Americas fromthe cordillera westward to the Pacific coast(Hershkovitz 1991a, 1993). Genera included in thisgroup are Calandrinia Kunth, Calyptridium Nutt.,Cistanthe Spach, Claytonia L., Lenzia Phil., LewisiaPursh, Lewisiopsis Govaerts (= Cistanthe sect.Strophiolum Hershk.), Montia L., and MontiopsisKuntze. Monophyly of the group remains unresolvedand depends upon the relations of Phemeranthus Raf.,distributed mainly in North America from the cordilleraeastward, and the Australian calandrinias (ParakeelyaHershk.; these species should be classified inRumicastrum Ulbr. according to J. G. West, pers. comm.,cf. Carolin 1987, 1993). However, molecular evidenceindicates that these taxa together form a well-supportedclade within the portulacaceous alliance (Hershkovitz& Zimmer 1997, 2000).

Species diversity of western AmericanPortulacaceae is divided approximately equallybetween North America and South America, andmuch of it is concentrated in California and northernChile. Notwithstanding several amphitropicaldisjunctions, most of the genera can be characterizedas predominantly or completely North American orSouth American. However, the present data indicatethat the largely South American Cistanthe sect.Amarantoides (Reiche) Carolin ex Hershk. andCistanthe sect. Philippiamra (Kuntze) Hershk. aremore closely related to the North AmericanCalyptridium than to the largely South AmericanCistanthe sect. Cistanthe. The data suggest thatthe first two sections may be preferably classified inCalyptridium. However, this and other nomenclaturalactions supported by the present analyses will bedeferred pending revision of type material.

This work presents analysis of diversity of bothITS and ycf3-trnS spacer sequences. ITS, a typically600 base-pair sequence, is the most widely appliedmolecular marker in interspecific phylogeneticanalyses in angiosperms (Hershkovitz et al. 1999).The ycf3-trnS intergenic spacer separates an openreading frame of unknown function (ycf3) from thechloroplast transfer RNA for serine bearing theanticodon GGA (trnS-GGA). These sequences havebeen determined for ca. 275 samples of western

American Portulacaceae, ca. 145 of which representSouth American (mostly Chilean) samples. Thepresent analyses incorporate 183 of these samples,emphasizing the South American samples butincluding sufficient samples from outside this regionto provide appropriate phylogenetic reference.Superficial examination of some of the gene treessuggests evolutionary radiation too rapid toreconstruct cladistically. This would be a reasonablescenario given evidence of relatively recent, rapid,and catastrophic events that shaped modern Chileanhabitats (Arroyo et al. 1988; Villagrán 1995). Moredetailed examination of the data in light of fieldobservations, however, reveals evidence for analternative scenario, in which the gene trees reflectnot rapid diversification, but effective homo-genization of the marker sequences via gene flow,thus obscuring a pattern of ecological andmorphological specialization that occurred over alonger historical period. This paper presentsevidence for both of these models and consequentimplications for diversification of the Chilean flora.

MATERIALS AND METHODS

PLANT MATERIALS

Table I provides collection data for the analyzedspecimens. For field collections, above-ground parts(variously, leaves, stems, flower buds, but not openflowers) from one or a few identical individuals, freefrom insect or other apparent damage or infection,were washed thoroughly with tap water and dried inclean paper towels prior to silica preservation.Herbarium specimens were prepared at the same time.Floristic, monographic, and other taxonomicreferences (see below) and museum specimens wereused to identify the materials. For a variety of reasons,identification of many of the Chilean samples ofCistanthe was difficult and in many cases notsuccessful: (1) of all the South American taxa, onlyMontiopsis subg. Montiopsis has beenmonographed (Ford 1992); (2) the most recentcomprehensive floristic revision of the Chilean taxais that of Reiche (1898, 1905); this treatment notesnumerous taxonomic problems and, nonetheless, itsreliability is questionable given the degree ofdiscordance with the monograph of Ford (1992); (3)succulent members of Cistanthe preserve poorly as

Andean region Portulacaceae Evolution: HERSHKOVITZ, M.

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herbarium specimens; (4) type specimens of manyCistanthe species are missing at SGO, and many ofthose present are too poorly preserved to aid inidentification; (5) several types of Cistanthe speciesrepresent populations now extinct in areas laterurbanized (and current observations of this groupindicate that localized populations aremorphologically distinctive); (6) as noted by Ford-Werntz & Peralta (2002), infrequency and irregularityof rain in the north of Chile hampers efforts to relocatepopulations from which taxa were described; and (7)notwithstanding the above, several collections frombetween 1998-2002 do not appear to correspond withany described taxa. The difficulty in identifying thespecies of the succulent taxa has been noted byHershkovitz (1991b) and Ford-Werntz & Peralta(2002). The latter synopsis (without key) of Cistantheincludes 22 species of Chile/Argentina/Peru but alsoa list of names of 28 taxa of unresolved application“where more studies as to relationship andcircumscription are necessary before the appropriatecombinations or synonymizations can be published”.In fact, as noted in the discussion, the historicaltaxonomic difficulties and present inability to identifynumerous specimens can be interpreted as anexpectation of the diversification scenariosproposed.

Eighteen of the DNA samples are not associatedwith voucher specimens. Data associated with suchspecimens should be interpreted circumspectfully,but not dogmatically. Several vouchers for DNAsamples published on previously (Hershkovitz &Zimmer 1997, 2000) appear to have been accidentallydiscarded. Other DNA samples represent cultivatedmaterial that was accessioned but not vouchered.The unvouchered materials are principally those ofthe outgroup and North American taxa and are henceperipheral to the theme of the present work. In anycase, all samples in the present work have somedegree of documentation verifying their source andidentity. Furthermore, I have handled and determinednearly all of the plant materials personally and haveperformed all of the DNA extractions.

MOLECULAR METHODS

DNA was extracted using the DNeasy Mini protocol(Quiagen, Inc.) and quantified using agarose gels.In some cases, additional purification was necessaryand accomplished via either PEG precipitation or by

a silica suspension (Boyle & Lew 1995). The PEGprotocol used equal volumes of sample and a fresh(< 2 weeks at 4C) solution of 20% PEG 8000 (Sigma)/2.5M NaCl. Following thorough mixing, the solutionis maintained at 37 °ðC for ca. 20 min, followed byhigh-speed centrifugation, careful (and thorough)removal of the supernatant, three washes of the pelletwith ca. 80% EtOH, drying, and resuspension in wateror AE buffer from the DNeasy kit (Quiagen). Tominimize the risk of cross contamination, especiallyby PCR products, extraction supplies and reagents,as well as PCR reagents, were maintained separatelyfrom those of post-PCR procedures. In addition, allsurfaces (including equipment, pipettors, and reagentcontainers) contacted during extraction and PCRreaction preparation were thoroughly washed with10% chlorine bleach solution prior to performingthese procedures. In several cases, DNA wasextracted from the herbarium specimencorresponding to the previously-extracted silicaspecimen in order to confirm sequencing results(Table I).

Amplification of ITS for sequencing generallyfollowed Hershkovitz and Zimmer (2000), i.e., double-strand amplification followed by separate asymmetricamplification of each strand using primers internalto the first. The amplification and sequencing primersare listed in Table 2. The amplification products weresequenced according to the Big-Dye Dye Terminator(Applied Biosystems) or DYEnamic ET Terminator(Pharmacia) cycle sequencing protocols. Sequencingreactions were electrophoresed on a 3100 GeneAnalyzer (Applied Biosystems). Most specimenswere sequenced in both directions. Exceptions weremade for chromatograms that were especially cleanin a single direction and which matched completelythose of samples sequenced in both directions.Many sequences of Cistanthe samples weresequenced in essentially one direction, because aninternal poly-T microsatellite-like repeat obliteratedclarity of the downstream signal.

PHYLOGENETIC ANALYSIS

Phylogenetic analyses were performed using PAUP4.0 (Swofford 2002) version b10. Analyses wereundertaken at the level of western AmericanPortulacaceae and at the level of each of the SouthAmerican clades. ITS and ycf3-trnS sequences wereanalyzed separately and in combination. The

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intergeneric analysis was performed as an update ofHershkovitz & Zimmer (2000) with additional criticalsamples and a cpDNA sequence. This analysisincluded representative sequences of the SouthAmerican genera analyzed in greater detail at theinfrageneric level and fewer representativesequences of the North American genera analyzedin greater detail elsewhere (Hershkovitz, submittedms.). This analysis provides an outgroup perspectiverelative to each of the primarily South Americangenera. Alignment was optimized manually.Unambiguous gaps were treated as independentcharacters. Up to four unordered states werepermitted for superposed (multistate) gaps. Ingeneral, when more than four gap states occurred,the alignment was ambiguous. Length variableregions that could not be aligned unambiguouslywere scored as unknown in all sequences or at leastin the sequences that could not be unambiguouslyaligned. Likewise, substitutions in length variableregions were scored as substitutions among thealignable sequences and as unknown in theunalignable sequences. Unweighted parsimony andassociated bootstrap (500 replicates) were appliedwith stepwise addition and tree bisection-reconnection. Bootstrap replicates held ten trees ateach addition step with maxtrees at 100 trees perreplicate. Partition homogeneity tests (500 replicates,maxtrees = 100) were performed to evaluate conflictin the two data sets. To visualize the total conflict inthe combined data, split decomposition wasperformed using SplitsTree 2.4 (Huson 1997) using«p» distances and refining for the maximum numberof possible quartets.

Maximum likelihood and minimum evolutionmethods were considered unnecessary for thepresent analyses. Previous analyses involving thesetaxa (Hershkovitz & Zimmer 1997, 2000) found thatall methods yielded largely unresolved intergeneric-level relations. At the infrageneric level, both thenumber of sequences and substitutions are very low,so that the variance in estimates of Markov modelsubstitution parameters used in distance andmaximum likelihood analysis would be very high(e.g., Posada 2001). Likewise, the short branchlengths at the infrageneric level and separatingseveral poorly resolved intergeneric nodes canmislead current implementations of Bayesianlikelihood methods (Suzuki et al. 2002; Cummings etal. 2003; P. O. Lewis, oral comm., 2003). In addition,

practical implementations of distance and likelihoodanalysis cannot take into account length variationcharacters, which provide a considerable proportionof the information, especially in the ycf3-trnSsequences. Finally, statistical inconsistency thatresults from using inaccurate substitution models isexpected mainly when divergence is high, e.g., whenthere are long branches.

RESULTS

ITS AND YCF3-TRNS SEQUENCE CHARACTERISTICS

Variability of the ITS region and ycf3-trnS spacer arecompared in Table III. Infra-and intergeneric ITSsequence divergences were calculated byHershkovitz & Zimmer (2000) this table is notreproduced here. Corrected intergeneric ITSsequence divergence (using ML distances as inHershkovitz & Zimmer 2000) among the SouthAmerican genera is generally on the order of 5%,except for that of Calandrinia sect. Calandrinia.Divergences of the latter to the other genera rangesfrom 10-13%. Maximum ITS divergence within therelevant taxa are on the order of: 4% (Calandriniasect. Calandrinia); 2% (Calandrinia sect. AcaulesReiche); 1% (Cistanthe sect. Cistanthe, Montiopsissubg. Montiopsis and Montiopsis subg.Dianthoideae (Reiche) D.I.Ford); and 0.5%(Cistanthe sections Amarantoides andPhilippiamra). Thus, net rates from a commonancestor are especially higher in the first taxon andespecially low in the last. Divergences within taxa ofthe ycf3-trnS sequences are approximately inproportion to the number of variable sites relative toITS.

As noted previously (Hershkovitz & Zimmer2000), the variable regions of ITS in western AmericanPortulacaceae are GC-rich except for Claytonia andMontia, in which the base bias may be absent or AT-rich. The ycf3-trnS sequences, as is typical of cpDNAspacers in general, are AT-rich, and include numerouspoly-A, poly-T, and poly-AT repeats 6-20 bases inlength. However, the aligned parsimony-informativesites of the ycf3-trnS sequences are not AT-, butrather GC-rich: 50-60% in these exemplars and 54%on average. Relative to ITS, variation in the ycf3-trnS sequences is more frequently in length ratherthan base substitution. At the intergeneric level, the


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ycf3-trnS spacer sequences include one region, ca.150 bases downstream of the 3’ end and 0-50 baseslong, that is unalignable between genera, althoughlargely alignable within genera. At the infragenericlevel, the length differences tend to be more easilyaligned than in ITS. The length differences commonlycorrespond to discrete direct repeats of three or morebases. In some cases, length variation occurs in apoly-A and/or –T region, resembling polymorphicmicrosatellite loci. The majority of the repeat variationobserved involved two states or two common statesand up to two additional rare states. A poly-T repeatvarying in length from 11 to 17 bases occurs in theycf3-trnS sequences of samples of Cistanthe sect.Cistanthe. Because of the high variability anddifficulty determining the exact number of Ts in thelonger repeats, this region was not included in thephylogenetic analysis.

INTERGENERIC-LEVEL ANALYSIS

Figures 1 and 2 show sample parsimony trees andparsimony bootstrap values for the ITS and ycf3-trnS data, respectively. Figure 3 shows the parsimonybootstrap consensus for the combined data. TheITS tree is considerably longer, indicating an overallhigher rate of evolution. However, the ITS tree alsohas much lower consistency and retention indicesthan the ycf3-trnS data, i.e., much of the lengthdifference is in apparently homoplasious changes.The ycf3-trnS data actually yield a larger number ofbranches with higher bootstrap proportions. Theoverall number of branches with higher bootstrapproportions is greater in the combined analysis thanin either data set alone. However, individual conflictsbetween the two data sets can be discerned, asdiagnosed by a reduction of particular bootstrapproportions in the combined analysis versus eitherof the separate analyses.

Circumscriptions of supraspecific taxa. BothITS and ycf3-trnS trees show a backbone structuresimilar to that in the previously published ITS-basedanalysis (Hershkovitz & Zimmer 2000). Monophylyof Cistanthe sensu Hershkovitz 1991a is stronglyrefuted by the combined data. Each tree shows >70% bootstrap proportions for the circumscriptionsof Cistanthe sect. Cistanthe, Lewisia, Claytonia,Montiopsis, Montiopsis subg. Montiopsis,Calandrinia, and Parakeelya. In one or the othertree, bootstrap proportions are lower, in some cases

< 50%, for the circumscriptions of Calyptridium,Cistanthe sections Amarantoides plusPhilippiamra, Montia, and Montiopsis subg.Dianthoideae. However, bootstrap proportions forthe combined data are > 78% for all of these clades.Intergeneric relations. Addition of critical samplesand the chloroplast data provide support forintergeneric relations not well-supported in theprevious analyses (Hershkovitz & Zimmer 1997,2000). The combined data show reasonably strongsupport for the existence of a “North American clade”comprising Lewisiopsis, Lewisia, Claytonia, andMontia. Support for this clade is weak in the separatedata analyses. The ycf3-trnS data support a sisterrelation between Lewisia and Lewisiopsis, butsupport is dramatically less in the combined analysis.In both data sets, the branch length from thehypothetical North American clade ancestor toLewisiopsis is much shorter than those of the othertaxa. Combined data support is also high for a cladecomprising Lenzia, Calyptridium, and Cistanthesections Amarantoides and Philippiamra. Again,support is weak in the separate data sets. Therelations of Parakeelya conflict in the parsimonyconsensus trees of the ITS and ycf3-trnS sequences(not shown). In the former, the genus is sister toCalandrinia, and in the latter to the North Americaclade. Oddly enough, neither position is supportedin the majority-rule bootstrap consensus of eitherdata set. In fact, the ycf3-trnS of the bootstrapmajority–rule tree (not shown) conflicts with theparsimony consensus in showing the strictlywestern American Portulacaceae as monophyletic,as in the combined data bootstrap, though withinsignificant support (51%).

RELATIONS AMONG SPECIES OF THE PREDOMINANTLY

SOUTH AMERICAN CLADES

1. Calandrinia. Parsimony trees for separate andcombined data analyses are presented in Figures 4-6. A splits graph is shown in Figure 7. Two sampleswere polymorphic at single sites in the ITS. Thegenotypes are indicated by “a” and “b”. These treescannot be rooted confidently by an outgroupcriterion, because the outgroups are approximatelyequally diverged from the Calandrinia species(Figures 1-2), and different outgroups root theingroup differently. The midpoint roots of the ITSand ycf3-trnS trees are different. The trees are

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arbitrarily rooted between the annuals (Calandriniasections Calandrinia and Monocosmia) andperennials (Calandrinia sect. Acaules), although thecombined data analysis per se supports a partitionbetween these groups. The ITS and ycf3-trnS trees areincongruent at certain points, but complementary atothers, as evidenced by the much larger number ofbranches with higher bootstrap proportions in thecombined versus the separate analysis. Totalincongruence can be visualized with the splits graph.The partition homogeneity test indicates that the twodata sets conflict (p = 0.001). Divergence among theperennials is uniformly low. Some interspecificdivergences among annuals are higher than others.Divergences between nearest neighbors are generallyproportional between ITS and ycf3-trnS, except forCalandrinia monandra DC. which shows muchhigher ITS than ycf3-trnS divergence.

The combined analysis supports three interspecificgroups: (1) C. menziesii (Hook.) Torr. & A. Gray and C.breweri S. Watson; (2) Calandrinia ciliata (Ruiz &Pav.) DC. and Calandrinia compressa Schrad. ex DC.;and (3) Calandrinia axilliflora Barnéoud and theunidentified collections from the Falkland Islands (IslasMalvinas). The first group is temperate NorthAmerican, although Calandrinia menziesii isintroduced in the southern hemisphere. The secondgroup is Central American and South American, andthe last is temperate South American. Support for thethird group is from the ITS data. The ycf3-trnSsequences support a different relationship for theFalklands plants. This incongruence is evident in thesplits graph. The ITS of one collection of Calandriniabreweri is different from the remainder and slightlymore similar to those of Calandrinia ciliata andCalandrinia compressa. Neither data set supports aclose relationship of Calandrinia monandra to anyparticular annual taxon. Sequence divergence amongthe perennial species is less than among the annualsand interspecific relations are mostly poorly supported.The morphologically diagnosed hybrid Calandriniacaespitosa x compacta has both the ITS and ycf3-trnSsequences of the sympatric Calandrinia compactaBarnéoud samples, but the two Calandrinia compactasamples are divergent in both sequences. The splitsgraph shows considerable reticulation among theCalandrinia compacta and Calandrinia caespitosaGillies ex Arn. samples.

2. Cistanthe sections Amarantoides and

Philippiamra. The separate and combined ITS andycf3-trnS trees are shown in Figures 8-10. Support formonophyly of the two sections together is sensitiveto sampling and sequence and depends on therelationships of Cistanthe ambigua (S.Watson)Carolin ex Hershk. Support for monophyly of the twosections minus Cistanthe ambigua is strongest (98%bootstrap support) in the combined data infrageneric-level analysis and weakest in the ITS intergeneric-levelanalysis (less than 50% bootstrap support). Supportfor monophyly of Calyptridium is strongest in thecombined data intergeneric-level analysis (80%bootstrap support) and weakest in both of the ycf3-trnS analyses (less than 50% bootstrap support). Thebootstraps do not support the inclusion of Cistantheambigua in either section. The trees are outgroup-rooted with Lenzia (cf. Figs. 1-2). Relations amongspecies of Calyptridium will be considered in moredetail elsewhere (Hershkovitz, in prep). The divergenceamong species of Cistanthe sections Amarantoidesand Philippiamra is extremely low, hence themonophyly of each section is neither supported norstrongly refuted. However, the ycf3-trnS sequences ofCistanthe (Philippiamra) amarantoides (Phil.) Carolinex Hershk. and most samples of Cistanthe(Amarantoides) calycina (Phil.) Carolin ex Hershk. areidentical and share at least two differences from allother samples.

3. Cistanthe sect. Cistanthe3a. Grandiflora group (Calandrinia sect. Cistanthesensu Reiche 1897). Figures 11-12 show the MP treesfor the ITS and ycf3-trnS data. Figure 13 shows thesplits graph for the combined data. The combined databootstrap is not shown because it includes only asingle partition, that separating Cistanthe sp. indet.,aff. Calandrinia crassifolia Phil. and Cistanthe sp.00-51 from the remaining taxa (84% support). Both treesare arbitrarily midpoint rooted, but the midpoint in theITS tree is among the many equally parsimonious rootspossible using the Rosulatae group as the outgroup.As is evident, divergence of both sequences is lowoverall, and many samples have identical sequences.The ITS data do not yield substantial bootstrapsupport for any relationships. Even though theinformative variation is much less, the ycf3-trnS dataproduce one reasonably strong bootstrap partition.This partition is absent, however, in the combined databootstrap consensus, whereas the combined datashow stronger support for a partition less strongly


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supported in each of the separate data sets. The lackof shared agreement between the two data sets isindicated by the reduced rescaled consistency indexin the combined versus separate data analyses, asignificant score in the partition homogeneity test (p =0.03) notwithstanding the low variation, and thereticulate appearance of the splits graph.

3b. Rosulatae group (= Calandrinia sectionsAndinae, Arenarie and Rosulatae sensu Reiche 1897).Figures 14-16 show the MP trees for the ITS and ycf3-trnS data. The ITS tree is arbitrarily midpoint rooted,but the root shown is among many equallyparsimonious roots possible using the Grandifloragroup sequences as outgroups. The midpoint root ofthe ITS data is the unequivocal root of the ycf3-trnSdata using outgroup rooting. However, this is not thesame root produced by midpoint rooting of the ycf3-trnS data. As with the Grandiflora group, variation islow for both sequences, and only the ycf3-trnS dataproduce any relatively high bootstrap proportions.Likewise, the combined data show generally reducedsupport for partitions in the separate data sets.However, conflict appears to be less than in theGrandiflora group, as indicated by the partitionhomogeneity test (p = 0.17). The splits graph (notshown) shows only one internal quadrangle. In threecases, support is increased in the combined datarelative to the separate data. One case is trivial, thatinvolving Cistanthe sp. indet., aff. Calandriniaoblongifolia Barnéoud and its putative hybridoffspring, which have identical ITS and ycf3-trnSsequences. Two other cases involve increased supportfor intertaxon relations between Cistanthe sp. indet.,aff. Calandrinia thyrsoidea Reiche, Cistanthecephalaphora (I.M.Johnst.) Carolin ex Hershk., andCistanthe maritima (Nutt. ex Torr. & A.Gray) Carolinex Hershk., and between Cistanthe sp. indet., aff.Calandrinia chamissoi Barnéoud and a sample ofCistanthe arenaria (Cham.) Carolin ex Hershk.However, Cistanthe arenaria appears as polyphyleticin both trees. Likewise, the samples of Cistanthelongiscapa (Barnéoud) Carolin ex Hershk. appear aspolyphyletic, and one sample of Cistanthe cymosa(Phil.) Carolin ex Hershk. (from near the type locality)shares the same ITS sequence as a sample of Cistanthelongiscapa collected at the same locality.

4. Montiopsis. ITS and ycf3-trnS parsimony treesare illustrated in Figures 17-19. Each subgenus is

outgroup-rooted with the other, although it isconceivable that high divergence between thesubgenera could produce rooting artifacts withinthem. In fact, the ITS and ycf3-trnS trees conflictin the placement of the root of each subgenus.However, the two data sets conflict in numerousaspects, evidenced in part by the reduction ofbootstrap proportions in the combined versusseparate analysis. In addition, the conflict isdemonstrated by the partition homogeneity test(p = 0.004).

4a. Montiopsis subgenus Dianthoideae. The ITSand ycf3-trnS trees conflict in their placement of thetaxa. The ycf3-trnS sequences of the Montiopsisgayana (Barnéoud) D.I.Ford samples are closelyrelated to those of Montiopsis sp. indet., aff.Calandrinia tricolor Phil., but the ITS sequencesare more divergent. The combined data consensusfavors the ycf3-trnS result.

4b. Montiopsis subgenus Montiopsis. The samplesof Montiopsis x trifida, Montiopsis uspallatensis(Phil.) D.I.Ford, and one of the samples of Montiopsistrifida (Hook. & Arn.) D.I.Ford were polymorphicfor ITS, and the last of these was also polymorphicfor ycf3-trnS. The genotypes are indicated withletters, as above. The polymorphic samples ofMontiopsis x trifida and Montiopsis trifida were eachpolymorphic for two positions in the ITS. Thiscreates four possible genotypes, of which twopossible extremes (different at both positions) werearbitrarily designated for analysis. Without cloning,it cannot be determined which sequences actuallyexist. Sequence divergence, especially of the ITS, islow relative to the number of taxa. Nonetheless, thecombined data suggests that the two data setsconflict. This conflict is less evident in the splitsgraph (not shown). Of the two clades supported by81% bootstrap support in the ycf3-trnS data, onlyone is present in the combined data bootstrapconsensus, and the support is lower. The sampleMontiopsis x trifida has both the ITS and ycf3-trnSsequence of Montiopsis trifida. One sample ofMontiopsis parviflora (Phil.) D.I.Ford has the ycf3-trnS sequence of Montiopsis trifida but the ITSsequence of the other Montiopsis parviflorasamples. However, the ITS divergence between theMontiopsis trifida and Montiopsis parviflorasamples is as little as one change.

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FIGURE 1. Most parsimonious (MP) tree for ITS data for western American Portulacaceae. Sample names are defined inTable I. One of 7146 trees, length 479. Rescaled consistency index (RC) 0.52, retention index (RI) 0.80. The tree isoutgroup rooted with samples of Phemeranthus and Talinum (cf. Hershkovitz & Zimmer 1997, 2000). Bootstrapproportions (BP; 500 replicates) greater than 50% are indicated. Two BP values less than 50% are indicated with anasterisk: these branches are in conflict with branches in the MP strict consensus. Note the longer branch lengthsassociated with samples of Claytonia and Montia and, to a lesser degree, Lewisia. The bootstrap consensus includes 13branches with BP values >70%.

FIGURA 1. Arbol más parsimonioso (MP) de los datos de ITS de Portulacaceae de América occidental. Los nombres demuestras se presentan en la Tabla I. Uno de 7146 árboles, largo 479, índice de consistencia rescalado (RC) 0,52, índice deretención (RI) 0,80. El árbol está enraizado con las muestras de Phemeranthus y Talinum (cf. Hershkovitz & Zimmer1997, 2000). Se indican las proporciones de “bootstrap” (BP, 500 réplicas) mayores que 50%. Se indican con un asteriscodos valores BP menores que 50%: estas ramas están en conflicto con las ramas en el árbol de MP del consenso estricto.Nótese las ramas largas asociadas con muestras de Claytonia y Montia y, en menor grado, Lewisia. El consenso debootstrap incluye 13 ramas con valores de BP >70%.

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ITSITSITSITSITS


21

FIGURE 2. Most parsimonious (MP) tree for ycf3-trnS data for western American Portulacaceae. Sample names are defined inTable I. One of 120 trees, length 384, RC 0.71, RI 0.88. The tree is outgroup rooted as in Fig. 1. BP values (500 replicates)greater than 50% are indicated. Two BP values less than 50% are indicated with an asterisk: these branches are in conflict withbranches in the MP strict consensus, Western American Portulacaceae (all taxa minus the outgroups and Parakeelya) arepartitioned in the bootstrap consensus (BP = 51%). Note the longer branch lengths associated with samples of Claytonia,Montia, and Lewisia (cf. Fig. 1). Note also longer branches associated with samples of Montiopsis (cf. Fig. 1). The bootstrapconsensus includes 19 branches with BP values >70%.

FIGURA 2. Arbol más parsimonioso (MP) para los datos de ycf3-trnS de Portulacaceae de América occidental. Los nombres demuestras se presentan en la Tabla I. Uno de 120 árboles, de largo 384, RC 0,71 y RI 0,88. El árbol está enraizado como en laFigura 1. Se indican valores de BP (500 réplicas) mayor que 50%. Se indican con un asterisco dos valores BP menores que50%: estas ramas están en conflicto con ramas en el árbol de MP del consenso estricto. Se separa en el consenso de bootstrap(BP = 51%) Portulacaceae de América occidental (todos los taxa menos los grupos externos y Parakeelya). Nótese las ramaslargas asociadas con muestras de Claytonia, Montia y Lewisia. Nótese también las ramas largas asociadas con muestras deMontiopsis (cf. Figura 1). El consenso de bootstrap incluye 19 ramas con valores de BP >70%.

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ycf3-trnSycf3-trnSycf3-trnSycf3-trnSycf3-trnS

Gayana Bot. 63(1), 2006

22

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FIGURE 3. Bootstrap consensus for combined ITS and ycf3-trnS data for western American Portulacaceae. Samplenames are defined in Table I. Analysis of the combined data yielded 104 MP trees, length 888, RC 0.58, RI 0.82. BPvalues (500 replicates) greater than 50% are indicated. The bootstrap consensus includes 25 branches with BP values>70%.

FIGURA 3. Consenso de bootstrap para los datos combinados de ITS y ycf3-trnS de Portulacaceae de América occidental.Los nombres de las muestras se presentan en la Tabla I. El análisis produjo 104 árboles de MP, de largo 888, RC 0,58y RI 0,82. Se indican valores de BP (500 réplicas) mayores que 50%. El consenso de bootstrap incluye 25 ramas convalores de BP >70%.

ITS + ycf3-trnSITS + ycf3-trnSITS + ycf3-trnSITS + ycf3-trnSITS + ycf3-trnS


23

FIGURE 4. MP tree for ITS data for Calandrinia. Sample names are defined in Table I. One of 101,179 trees, length 57,RC 0.91, RI 0.98. BP values (500 replicates) greater than 50% are indicated. Lower case letters following duplicatedsample names denote samples having more than one ITS genotype. The tree is rooted with samples of Calandriniasect. Acaules as outgroup. Midpoint rooting of this tree places the root between Calandrinia monandra (the longestbranch) and the remaining taxa. The bootstrap consensus includes 4 branches with BP values >70%.

FIGURA 4. Arbol más parsimonioso (MP) para los datos de ITS de Calandrinia. Los nombres de muestras se presentanen la Tabla I. Uno de 101.179 árboles, de largo 57, RC 0,91 y RI 0,98. Se indican los valores de BP (500 réplicas)mayores que 50%. Las letras en minúsculas que acompañan los nombres de muestras corresponden a muestras con másde un genotipo de ITS. El árbol se enraíza con las muestras de Calandrinia sect. Acaules como grupo externo. Si seenraíza el árbol por el criterio de “midpoint”, la raíz se encuentra entre Calandrinia monandra (la rama más larga) y elremanente de los taxa. El consenso de bootstrap incluye 4 ramas con valores de BP >70%.

Calandrina - ITS

Gayana Bot. 63(1), 2006

24

FIGURE 5. MP tree for ycf3-trnS data for Calandrinia. Sample names are defined in Table I. One of 2 trees, length 41, RC1.0, RI 1.0. BP values (500 replicates) greater than 50% are indicated. Lower case letters following duplicated samplenames denote samples having more than one ITS genotype (cf. Fig. 4). The tree is rooted with samples of Calandriniasect. Acaules as outgroup. The bootstrap consensus includes 6 branches with BP values >70%.

FIGURA 5. Arbol más parsimonioso (MP) para los datos de ycf3-trnS de Calandrinia. Los nombres de las muestras sepresentan en la Tabla I. Uno de 2 árboles, de largo 41, RC 1,0 y RI 1,0. Se indican los valores de BP (500 réplicas)mayores que 50%. Las letras en minúsculas que acompañan los nombres de muestras corresponden a muestras con másde un genotipo de ITS (cf. Figura 4). El árbol se enraíza con las muestras de Calandrinia sect. Acaules como grupoexterno. El consenso de bootstrap incluye 6 ramas con valores de BP >70%.

Calandrinia - ycf3-trnS


25

FIGURE 6. Bootstrap consensus for combined ITS and ycf3-trnS data for Calandrinia. Sample names are defined in Table I. BPvalues (500 replicates) greater than 50% are indicated. Lower case letters following duplicated sample names denote sampleshaving more than one ITS genotype (cf. Figure 4). The tree is rooted with samples of Calandrinia sect. Acaules as outgroup.Analysis of the combined data yielded 19 MP trees, length 103, RC 0.89, RI 0.98. The bootstrap consensus includes 14branches with BP values >70%.

FIGURA 6. Consenso de bootstrap para los datos combinados de ITS y ycf3-trnS de Calandrinia. Los nombres de lasmuestras se presentan en la Tabla I. Se indican los valores de BP (500 réplicas) mayores que 50%. Las letras enminúsculas que acompañan los nombres de muestras corresponden a muestras con más de un genotipo de ITS (cf.Figura 4). El árbol se arraiga con las muestras de Calandrinia sect. Acaules como grupo externo. El análisis de los datoscombinados produjo 19 árboles de MP, de largo 103, RC 0,89 y RI 0,98. El consenso de bootstrap incluye 14 ramas convalores de BP >70%.

Calandrinia - ITS + ycf3-trnS

Gayana Bot. 63(1), 2006

26

FIGURE 7. Splitsgraph for combined ITS and ycf3-trnS data for Calandrinia. For simplicity, duplicate samples of the sametaxon identical for both sequences were omitted. The omitted samples can be deduced from Figures 4 and 5. The splitsanalysis used “p” distances and was refined over the maximum possible number of quartets.

FIGURA 7. Splitsgraph de los datos combinados de ITS y ycf3-trnS de Calandrinia. Para simplicidad, muestras duplicadasdel mismo taxón, e iguales para las dos secuencias fueron eliminadas. Las muestras eliminadas pueden ser deducidas desdelas Figuras 4 y 5. El análisis de “splits” incorporó distancias de “p” y fue refinado sobre el número máximo de cuartetosposibles.

Calandrinia - ITS + ycf3-trnS


27

FIGURE 8. MP tree for ITS data for Lenzia, Calyptridium and Cistanthe sections Amarantoides and Philippiamra.Sample names are defined in Table I. One of 16 trees, length 55, RC 0.83, RI 0.93. BP values (500 replicates) greaterthan 50% are indicated. The tree is outgroup-rooted using Lenzia (cf. Figs. 1-3). The bootstrap consensus includes 3branches with BP values >70%.

FIGURA 8. Arbol más parsimonioso para los datos de ITS de Lenzia, Calyptridium y Cistanthe secciones Amarantoidesy Philippiamra. Los nombres de las muestras se presentan en la Tabla I. Uno de 16 árboles, de largo 55, RC 0,83 y RI0,93. Se indican los valores de BP (500 réplicas) mayores que 50%. El árbol se enraíza con las muestras de Lenzia comogrupo externo (cf. Figuras 1-3). El consenso de bootstrap incluye 3 ramas con valores de BP >70%.

ITS

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FIGURE 9. MP tree for ycf3-trnS data for Lenzia, Calyptridium and Cistanthe sections Amarantoides and Philippiamra.Sample names are defined in Table I. One of 1 tree, length 37, RC 0.88, RI 0.96. BP values (500 replicates) greater than50% are indicated. The tree is outgroup-rooted using Lenzia (cf. Figs. 1-3). The bootstrap consensus includes 2branches with BP values >70%.

FIGURA 9. Arbol más parsimonioso para los datos de ycf3-trnS de Lenzia, Calyptridium y Cistanthe secciones Amarantoidesy Philippiamra. Los nombres de las muestras se presentan en la Tabla I. El árbol más parsimonioso, de largo 37, RC0,88 y RI 0,96. Se indican los valores de BP (500 réplicas) mayores que 50%. El árbol se enraíza con las muestras deLenzia como grupo externo (cf. Figuras 1-3). El consenso de bootstrap incluye 2 ramas con valores de BP >70%.

ycf3-trns


29

FIGURE 10. Bootstrap consensus for combined ITS and ycf3-trnS data for Lenzia, Calyptridium and Cistanthe sectionsAmarantoides and Philippiamra. Sample names are defined in Table I. BP values (500 replicates) greater than 50% areindicated. The tree is outgroup-rooted using Lenzia (cf. Figs. 1-3). Analysis of the combined data yielded 8 MP trees,length 94, RC 0.82, RI 0.93. The bootstrap consensus includes 3 branches with BP values >70%.

FIGURA 10. Consenso de bootstrap para los datos combinados de ITS y ycf3-trnS de Lenzia, Calyptridium y Cistanthesecciones Amarantoides y Philippiamra. Los nombres de las muestras se presentan en la Tabla I. Se indican los valoresde BP (500 réplicas) mayores que 50%. El árbol se enraíza con las muestras de Lenzia como grupo externo (cf. Figuras1-3). El consenso de bootstrap incluye 3 ramas con valores de BP >70%.

ITS + ycf3-trns

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FIGURE 11. MP tree for ITS data for Cistanthe sect. Cistanthe, Grandiflora group. Sample names are defined in Table I.Roman numerals refer to Chilean Region of origin; M refers to the Metropolitan Region (see Table I). One of 251 trees,length 16, RC 0.78, RI 0.89. BP values (500 replicates) greater than 50% are indicated. The tree is midpoint-rooted.

FIGURA 11. Arbol más parsimonioso para los datos de ITS de Cistanthe sect. Cistanthe, grupo Grandiflora. Los nombresde las muestras se presentan en la Tabla I. Los números romanos se refieren a la región de Chile; M se refiere a la RegiónMetropolitana (ver Tabla I). Uno de 251 árboles, de largo 16, RC 0,78 y RI 0,89. Se indican los valores de BP (500réplicas) mayores que 50%. El árbol se enraíza con el criterio de “midpoint”.

Cistanthe sect. CistantheGrandiflora group

ITS


31

FIGURE 12. MP tree for ycf3-trnS data for Cistanthe sect. Cistanthe, Grandiflora group. Sample names are defined in Table I.Roman numerals refer to Chilean Region of origin; M refers to the Metropolitan Region (see Table I). Single MP tree,length 5, RC 1.0, RI 1.0. BP values (500 replicates) greater than 50% are indicated. The tree is midpoint-rooted.

FIGURA 12. Arbol más parsimonioso para los datos de ycf3-trnS de Cistanthe sect. Cistanthe, grupo Grandiflora. Losnombres de las muestras se presentan en la Tabla I. Los números romanos se refieren a la región de Chile; M se refiere ala Región Metropolitana (ver Tabla I). El único árbol más parsimonioso, de largo 5, RC 1,0 y RI 1,0. Se indican los valoresde BP (500 réplicas) mayores que 50%. El árbol se enrraíza con el criterio de “midpoint”.


ycf3-trnS

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FIGURE 13. Splitsgraph for combined ITS and ycf3-trnS data for Cistanthe sect. Cistanthe, Grandiflora group. Samplenames are defined in Table I. Roman numerals refer to Chilean Region of origin; M refers to the Metropolitan Region (seeTable I). The splits analysis used “p” distances and was refined over the maximum possible number of quartets.

FIGURA 13. Splitsgraph para los datos combinados de ITS y ycf3-trnS de Cistanthe sect. Cistanthe, grupo Grandiflora. Losnombres de las muestras se presentan en la Tabla I. Los números romanos se refieren a la región de Chile; M se refiere ala Región Metropolitana (ver Tabla I). El análisis de “splits” incorporó distancias de “p” y fue refinado sobre el númeromáximo de cuartets posibles.


ycf3-trnS


33

FIGURE 14. MP tree for ITS data for Cistanthe sect. Cistanthe, Rosulatae group. Sample names are defined in Table I.Roman numerals refer to Chilean Region of origin; M refers to the Metropolitan Region; MENDOZA refers to MendozaProvince, Argentina; CAL refers to California, USA (see Table I). One of 5 trees, length 34, RC 0.84, RI 0.96. BP values(500 replicates) greater than 50% are indicated. Rooting is arbitrarily in agreement with Figure 15, but the midpoint rootoccurs along the basal split shown.

FIGURA 14. Arbol más parsimonioso para los datos de ITS de Cistanthe sect. Cistanthe, grupo Rosulatae. Los nombres delas muestras se presentan en la Tabla I. Los números romanos se refieren a la región de Chile; M se refiere a la RegiónMetropolitana; MENDOZA a la provincia de Mendoza, Argentina; CAL a California, EEUU (ver Tabla I). Uno de 5árboles, de largo 34, RC 0,84 y RI 0,96. Se indican los valores de BP (500 réplicas) mayores que 50%. El árbol se enraízaen una manera arbitraria en acuerdo con Figura 15, pero la raíz de midpoint se encuentra en la divergencia basal.

Cistanthe sect. CistantheRosulatae group

ITS

Gayana Bot. 63(1), 2006

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FIGURE 15. MP tree for ycf3-trnS data for Cistanthe sect. Cistanthe, grupo Rosulatae. Sample names are defined in Table I.Roman numerals refer to Chilean Region of origin; M refers to the Metropolitan Region; MENDOZA refers to MendozaProvince, Argentina; CAL refers to California (see Table I). Single MP tree, length 17, RC 0.93, RI 0.98. BP values (500replicates) greater than 50% are indicated. The root is based on outgroup rooting using the Cistanthe sect. Cistanthe,Grandiflora group ycf3-trnS data.

FIGURA 15. Arbol más parsimonioso para los datos de ycf3-trnS de Cistanthe sect. Cistanthe, grupo Rosulatae. Losnombres de las muestras se presentan en la Tabla I. Los números romanos se refieren a la región de Chile; M se refierea la Región Metropolitana; MENDOZA a la provincia de Mendoza, Argentina; CAL a California, EEUU (ver Tabla I).El árbol más parsimonioso, de largo 17, RC 0,93 y RI 0,98. Se indican los valores de BP (500 réplicas) mayores que50%. El árbol se enraíza con los datos de ycf3-trnS de Cistanthe sect. Cistanthe, grupo Grandiflora.

Cistanthe sect. CistantheRosulatae group

ycf3-trnS


35

FIGURE 16. Bootstrap consensus for combined ITS and ycf3-trnS data for Cistanthe sect. Cistanthe, Rosulatae group.Sample names are defined in Table I. Roman numerals refer to Chilean Region of origin; M refers to the MetropolitanRegion; MENDOZA refers to Mendoza Province, Argentina; CAL refers to California, USA (see Table I). BP values(500 replicates) greater than 50% are indicated. The tree is midpoint-rooted.

FIGURA 16. Consenso de bootstrap para los datos combinados de ITS y ycf3-trnS de Cistanthe sect. Cistanthe, grupoRosulatae. Los nombres de las muestras se presentan en la Tabla I. Los números romanos se refieren a la región de Chile;M se refiere a la Región Metropolitana; MENDOZA a la provincia de Mendoza, Argentina; CAL a California, EEUU(ver Tabla I). Se indican los valores de BP (500 réplicas) mayores que 50%. El árbol se enraíza con el criterio demidpoint.

Cistanthe sect. CistantheRosulatae groupITS + ycf3-trnS

Gayana Bot. 63(1), 2006

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FIGURE 17. MP tree for ITS data for Montiopsis. Sample names are defined in Table I. One of 2 trees, length 27, RC 1.0,RI 1.0. BP values (500 replicates) greater than 50% are indicated. Lower case letters following duplicated sample namesdenote samples having more than one ITS genotype. The tree is rooted between subgenera Montiopsis and Dianthoides.The bootstrap consensus includes 4 branches with BP values >70%.

FIGURA 17. Arbol más parsimonioso para los datos de ITS de Montiopsis. Los nombres de las muestras se presentan enla Tabla I. Uno de 2 árboles, de largo 27, RC 1,0 y RI 1,0. Se indican los valores de BP (500 réplicas) mayores que 50%.Las letras en minúsculas que acompañan los nombres de muestras corresponden a muestras con más de un genotipo deITS. El árbol se enraíza entre los subgéneros Montiopsis y Dianthoideae. El consenso de bootstrap incluye 4 ramas convalores de BP >70%.

MontiopsisITS


37

FIGURE 18. MP tree for ycf3-trnS data for Montiopsis. Sample names are defined in Table I. One of 174 trees, length 69,RC 0.93, RI 0.99. BP values (500 replicates) greater than 50% are indicated. Lower case letters following duplicatedsample names denote samples having more than one ITS genotype. The tree is rooted between subgenera Montiopsis andDianthoides. The bootstrap consensus includes 5 branches with BP values >70%.

FIGURA 18. Arbol más parsimonioso para los datos de ycf3-trnS de Montiopsis. Los nombres de las muestras se presentanen la Tabla I. Uno de 174 árboles, de largo 69, RC 0,93 y RI 0,99. Se indican los valores de BP (500 réplicas) mayores que50%. Las letras en minúsculas que acompañan los nombres de muestras corresponden a muestras con más de un genotipode ITS. El árbol se enraíza entre los subgéneros Montiopsis y Dianthoideae. El consenso de bootstrap incluye 5 ramas convalores de BP >70%.

Montiopsisycf3-trnS

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FIGURE 19. Bootstrap consensus for combined ITS and ycf3-trnS data for Montiopsis. Sample names are defined in Table I. BPvalues (500 replicates) greater than 50% are indicated. Lower case letters following duplicated sample names denote sampleshaving more than one ITS genotype. The tree is rooted between subgenera Montiopsis and Dianthoides. Analysis of thecombined data yielded > 1,000,000 MP trees, length 104, RC 0.86, RI 0.97. The bootstrap consensus includes 4 brancheswith BP values >70%.

FIGURA 19. Consenso de bootstrap para los datos combinados de ITS y ycf3-trnS de Montiopsis. Los nombres de lasmuestras se presentan en la Tabla I. Se indican los valores de BP (500 réplicas) mayores que 50%. Las letras en minúsculasque acompañan los nombres de muestras corresponden a muestras con más de un genotipo de ITS. El árbol se enraízaentre los subgéneros Montiopsis y Dianthoideae. Los análisis de los datos en combinación produjo > 1.000.000 árbolesmás parsimoniosos, de largo 104, RC 0,86 y RI 0,97. El consenso de bootstrap incluye 5 ramas con valores de BP >70%.

MontiopsisITS + ycf3-trnS


39

DISCUSSION

Comparison of its and ycf3-trnS sequence data inphylogenetic reconstruction of western americanportulacaceae

For western American Portulacaceae as a whole,informative variability of the ycf3-trnS spacer isapproximately 10% greater than that of ITS (cf. Smallet al. 1998). At the generic level, the variation isapproximately equal. These comparisons includeinformative variation in sequence length (indels), butthese characteristics must be considered a priori atleast as robust as substitutions, if not more so. Inparticular, assuming that the probability of insertionor deletion of particular sequence lengths is, at leastwithin the range of observed length differences, notsystematically biased, then the probability of aparticular length variation is related to the probabilityof all possible length variants. In contrast, asubstitution has three possible outcomes. Thus, thehigh information content of indels balances theirpossible disadvantages, e.g., lack of well-developedmethods of incorporating indels into stochasticmodels of sequence evolution. In parsimonyanalysis, the ycf3-trnS sequences also showed equalor, in the case of all western American Portulacaceae,substantially lower levels of homoplasy than ITS.Although the two sequences appear to besubstantially complementary in their phylogeneticsignal, several cases of conflict were noted. Cronnet al. (2002) also found that chloroplast sequencecharacters showed lower homoplasy than that ofvarious nuclear sequences, and that chloroplast andnuclear sequences yielded conflicting phylogeneticsignal. Interpretation of the conflict in this case wasproblematic. The authors doubted but could notreject that the conflict was the result of interspecificgene flow. They also noted that analysis of lengthvariation versus substitution in the chloroplastsequences also conflicted, but they were unable todetect evidence that substitutions in the chloroplastsequences were systematically biased in a way thatwould yield spurious interpretation ofsynapomorphies. In the present infrageneric-levelanalyses, the number of informative substitutionsand length variations was too low to evaluatestatistically their degree of consistency.

The ITS tree shows relatively longer branchesleading to the North American genera Claytonia,Montia, and less markedly, Lewisia. Longer branches

for these taxa also occur in the ycf3-trnS tree.Hershkovitz & Zimmer (2000) proposed that rates ofITS evolution are higher in these taxa, and that therate difference was related to a shift in basecomposition at the parsimony–informative sites froma strong GC to a strong AT bias. The ycf3-trnS datamight suggest that the rate increase in the ITS andchloroplast are also related. However, no comparableshift in base composition is apparent in the ycf3-trnS sequences: informative sites in the NorthAmerican taxa are 50-54% GC, generally belowaverage for the exemplars as a whole, but not lowerthan in samples from the other genera. In contrast tothe apparent correlation of rates for the NorthAmerican taxa, the branch lengths leading toMontiopsis are different in the two data sets, relativelyshort in the ITS data and relatively long in the ycf3-trnS data. In any case, differences in branch lengthsin either data set at the intergeneric level are notrelated to life form, i.e., a shift to the annual habit.The North American taxa are all perennial (Lewisia)or include both perennial and annual species(Claytonia and Montia). Likewise, species ofMontiopsis subg. Dianthoideae are all perennials,so it is not clear why their ycf3-trnS divergences aregreater than among the Rosulatae group of Cistanthesect. Cistanthe, which are mainly annuals. However,differences in divergence coincident with differencesin life form were evident within some of the genera(see below).

INTERGENERIC RELATIONS AMONG WESTERN AMERICAN

PORTULACACEAE

Unfortunately, increased ITS sampling and theaddition of a chloroplast marker did little to improveresolution of relations among genera beyond whatwas reported by Hershkovitz & Zimmer (2000). Theexplanation proposed previously was that the generaradiated explosively. An alternative possibility, viz.,the gene trees reflect artifacts of hybridization, isconsidered following discussion of the individualgenera. However, the new data do clarify relationsinvolving Lewisiopsis and Lenzia, as discussedbelow. Because of the poor resolution, the analysisprovides only limited justification for rooting of thevarious infrageneric trees, and no justification at allin the case of Calandrinia. The intergeneric treeappears to provide clear rooting points or outgroupsfor the Grandiflora and Rosulatae groups of

Gayana Bot. 63(1), 2006

40

Cistanthe sect. Cistanthe (each other), Montiopsis(between subgenera Montiopsis and Dianthoideae),and Calyptridium plus Cistanthe sectionsAmarantoides and Philippiamra (Lenzia), but theserootings may be misleading. Specifically, the branchlengths of the root branches are long relative to therespective ingroup branch lengths. This mayintroduce long-branch artifacts.

1. Lewisiopsis and the North American clade.Evidence for the North American clade is relevantto the understanding of the relation between theAndean and non-Andean taxa. Lewisiopsis tweedyi(A.Gray) Govaerts historically was classified inLewisia. Morphological analysis (Hershkovitz 1992)indicated that this species lacked synapomorphiesof Lewisia and, on the balance, was more similar tospecies of Cistanthe than to any species of Lewisia.For this reason, the species was transferred toCistanthe. In fact, that analysis indicated thatLewisiopsis tweedyi was not more similar to anyLewisia species than to Claytonia species. Theprevious ITS analysis (Hershkovitz & Zimmer 2000)provided no support for relations to Cistanthe andonly weak support for a “North American” cladecomprising Lewisiopsis tweedyi, Lewisia, and theClaytonia/Montia clade. However, the moleculardata were remarkably parallel to the morphologicalin that the ITS sequence of Lewisiopsis was highlyplesiomorphic relative to Lewisia and Claytonia/Montia. In fact, it is more similar to the Cistanthethan to the North American taxa sequences. Thepresent ITS analysis is in agreement. The ycf3-trnSsequence data provide relatively strong support fora sister-relation between Lewisiopsis and Lewisia,although, as with ITS, the ycf3-trnS sequence ofLewisiopsis tweedyi is also plesiomorphic. Supportfor the Lewisiopsis-Lewisia clade, however, issensitive to sampling in the other North Americangenera (data not shown). The analysis of thecombined data provide strong support for the NorthAmerican clade but weak support for theLewisiopsis-Lewisia clade. The reduction of supportfor the latter clade in the combined analysissuggests that the ITS and ycf3-trnS sequencesconflict somewhat. The relictuality of Lewisiopsisamong the North American taxa is discussed ingreater detail elsewhere (Hershkovitz, in prep.)

2. The relations of Lenzia. Surprising given its

morphology (cf. Hershkovitz 1993), the monotypicLenzia appears as the sister group to Calyptridiumplus Cistanthe sections Amarantoides andPhilippiamra. Lenzia chamaepitys Phil. is anacaulescent high alpine chaemophyte distributedbetween ca. 22-31°ð S. At ca. 30°ð S, it was observedto occur among the highest elevation plant species(4200 m). The plant is a 2-3 cm tall tuft of largelymembranous awl-shaped leaves. Reportedly, it hassolitary, axillary flowers (Carolin 1993). The plantsare connected by slender rhizomes (pers.observ.), acharacteristic not previously reported. Amongwestern American Portulacaceae, this acaulescent,perennial growth form is most suggestive ofCalandrinia sect. Acaules, especially thediminutive and rhizomatous Calandrinia compactathat occurs in the alpine zone in the same region.Cistanthe sections Amarantoides andPhilippiamra, in contrast, are all succulent,ramified lowland desert annuals with multiflorous(20-100 flowers) inflorescences. As noted inHershkovitz (1991a, c; see also Carolin 1987), theplants have the same vegetative andinflorescence morphology and leaf anatomy as inCistanthe. The flowers of Calyptridium plusCistanthe sections Amarantoides andPhilippiamra are smaller and more congested andthe sepals more membranous than in mostCistanthe species, but these characteristicsintergrade in the two. Calyptridium are alsomainly low-elevation annuals. The higher-elevation condition and perenniality appear to bederived in this genus (Hershkovitz in prep.). Theonly reported characteristic of Lenzia that iscoincident with Calyptridium plus Cistanthesections Amarantoides and Philippiamra ismembranous sepal texture (Reiche 1905). However,the leaves of Lenzia are also membranous. Thus,even in hindsight, morphology and ecologyprovide little if any hint of the sister relationbetween Lenzia and Calyptridium plus Cistanthesections Amarantoides and Philippiamra .Although the evidence for the relations of Lenziarefute the hypothesized monophyly of Cistanthesensu Hershkovitz 1991a, the molecular datasupport the previously hypothesized closerelationship between the North AmericanCalyptridium and South American Cistanthesections Amarantoides and Philippiamra(Carolin 1987; Hershkovitz 1993).


41

INFRAGENERIC RELATIONS AMONG WESTERN AMERICAN

PORTULACACEAE

1. Calandrinia. As noted, the rooting of theCalandrinia trees remains problematic, but the ITSand ycf3-trnS data are at least consistent with theclassification of Calandrinia in three sections. Inparticular, the annual and perennial taxa arepartitioned, and the data do not indicate a clearrelationship for Calandrinia sect. MonocosmiaHershk. As noted in Hershkovitz (1993), Calandrinia(Monocosmia) monandra resembles the annualsvegetatively, but its inflorescence and flowers aredistinctive, so much so that the species wastraditionally classified not only in the separate genusMonocosmia Fenzl, but in a different tribe ofPortulacaceae. The ITS and ycf3-trnS data conflictat certain points, which is consistent with otherinformation that hybridization may occur amongcertain taxa, as discussed below. The data alsocontribute to the taxonomic understanding ofCalandrinia menziesii, Calandrinia feltonii Skotts.,and two unidentified forms.

Calandrinia menziesii, which is distributed fromBaja California to British Columbia, historically hasbeen considered as a variety of the Central Americanand South American Calandrinia ciliata (e.g.,Macbride 1931; Munz & Keck 1968) or even as thesame species (e.g., Kelley 1993). Calandriniamenziesii also has been considered as a variety oras the same species as Calandrinia caulescensKunth (e.g., Gray 1887), but the latter is generallyconsidered a taxonomic synonym of Calandriniaciliata. In any case, there has never been a clearmorphological distinction between the temperateNorth American versus the Central American andSouth American plants. The characteristics ofCalandrinia menziesii mentioned by Macbride(1931), “more numerous stamens (often) and largerflower,” are actually highly polymorphic in thisspecies. In fact, the taxonomic recognition ofCalandrinia menziesii appears to have been amongNorth American populations rather than betweenthese and those of Central America and SouthAmerica (e.g., Gray 1887; Rydberg 1928). Both theITS and ycf3-trnS data show that the Calandriniamenziesii samples (spanning Baja California toBritish Columbia) are divergent from those ofCalandrinia ciliata (Mexico and Argentina).Ironically, the Calandrinia ciliata sequences are

identical to those of Calandrinia compressa, amorphologically distinctive species endemic tocentral/southern Chile, at least 1000 km disjunct fromCalandrinia ciliata. However, some herbariumcollections from northernmost Chile and in the rangeof Calandrinia cililata appear to be intermediate inmorphology between Calandrinia ciliata andCalandrinia compressa. Thus, the possibility ofhybridization involving Calandrinia compressashould be considered.

It is further ironic that evidence for hybridizationinvolving Calandrinia compressa emerges from thestudy of Calandrinia menziesii apparentlyintroduced to the Falkland Islands (Islas Malvinas),where it has been known as Calandrinia feltonii.Calandrinia feltonii had been presumed to be anendemic species that was extirpated in the wildbecause of grazing and other human activity(Davidson 1975; Woods 1994, 2000a). I had con-sidered this species to be the same as Calandriniaciliata (sensu Kelley 1993, including Calandriniamenziesii), as indicated in my 1992 annotation of thetype (Skottsberg 1910, UPS!, SGO!). The sameopinion was shared by Peralta (1999). Circumstantialevidence suggested that this plant may have beenintroduced to the Falkland Islands: the absence ofpopulations in undisturbed localities, its long historyof cultivation on the islands, the remoteness of theFalklands from the nearest wild populations ofCalandrinia ciliata sensu lato (in the highlands westof Buenos Aires), the introduction and subsequentestablishment of this species in Australia, NewZealand, and, most recently, India (Pande 2001), thefrequent cultivation of this species in Europeanbotanical gardens in the late 19th century, and thestrategic location of the Falkland Islands in theshipping route between Europe and the AmericanPacific coast. Nonetheless, Calandrinia feltoniibecame a keystone species of Falklands Islandsconservation efforts, even featured on aconservation-themed Falkland Islands postagestamp. The search for wild populations ofCalandrinia feltonii led to the discovery of anapparently distinct Calandrinia (Woods 2000b).Actually, the “new” form [e.g., R. W. Woods, s.n. 24Dec 2001 (Falkland Islands National Herbarium,photo!)] had been cultivated and collected as earlyas 1895 [L. H. Firman s.n. (K, received June 1895), R.Woods, pers. comm.], but the earlier collection waspresumed to be Calandrinia feltonii. From

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photographs (courtesy of R. Woods), theunidentified taxon more closely resembles theChilean endemic Calandrinia axilliflora, butnonetheless appears distinct. The present datashow that the Falkland Island specimens purportingto be Calandrinia feltonii have ITS and ycf3-trnSsequences identical to Calandrinia menziesii.Given the genetic distinction between Calandriniamenziesii and Calandrinia ciliata, this corroboratesthe hypothesis of introduction. The data for thepurportedly new form (C_sp_Mal1 and C_sp_Mal2)are conflicting. The ITS tree corroborates the closerelationship of this plant with Calandriniaaxilliflora of Chile, but the ycf3-trnS sequences aremore similar to those of Calandrinia ciliata andCalandrinia compressa. Although materials usedin the present study were not vouchered, theauthenticity of their identity is supportedcircumstantially by the DNA data. In particular,sequences of the samples purported to beCalandrinia feltonii conform as expected toCalandrinia menziesii, and ITS sequences of thesamples purported to be the form similar toCalandrinia axilliflora likewise conform to thisspecies. Moreover, the sequence data indicate thatthese samples may be hybrids.

Subsequent examination of herbarium materialhas revealed that plants resembling the unidentifiedFalkland Islands form have been collected in coastalsouthern Chile, mainly between Concepción andValdivia [e.g., Lechler 585 (P!), C. Gay 65 (P!)]. Apossible scenario supported by the sequence datais that the unidentified form is actually a hybridbetween Calandrinia axilliflora and eitherCalandrinia compressa or Calandrinia ciliata, andthat, like Calandrinia menziesii, it has beenintroduced into the Falkland Islands. Also consistentwith the data is the possibility that Calandriniacompressa itself is of hybrid origin. This wouldexplain why this ecologically and morphologicallydistinctive species shares genotypes withCalandrinia ciliata.

As noted in Table I, another unidentifiedcalandrinia, Hershkovitz 99-22B, occurs among theperennial species, described and illustrated inHoffmann et al. (1998, pp. 56-57). This plant is knownonly from two small populations (< one hectare) in aski area east of Santiago. It has a jointed pedunclecharacteristic of (sub)tropical perennial calandriniasthat occur only north of the arid diagonal, a virtually

plantless region that transects the Andes at ca. 24°SS. This natural phytogeographic barrier separatesthe southern Andes, which receives moisture inwinter via an Antarctic polar front, from the centralAndes, which receives moisture in the summermonths via the Intertropical Convergence (Arroyoet al. 1988). Relatively few species (as currentlyrecognized) are distributed on both sides of thediagonal (Arroyo et al. 1988, unpublished data), andamong Calandrinia species, only Calandriniacompacta, distributed from Bolivia to central Chileand Argentina (Peralta 1999). The presence of theunidentified form in central Chile could be explainedby numerous scenarios involving, variously,dispersal, relictuality, human influence, and/orhomoplasy. ITS and ycf3-trnS sequence divergencesare low among the perennials, hence insufficient todefinitively corroborate one hypothesis overanother. Nonetheless, the ITS sequence of theunidentified form is more similar to those of centralChilean species while the ycf3-trnS sequence is moresimilar to that of the northern species, Calandriniaacaulis Kunth. This introduces the possibility thatthe unidentified form represents a hybrid.

The sampling also includes a naturalmorphologically intermediate hybrid betweenCalandrinia compacta and Calandrinia caespitosathat was collected where the two species weregrowing in intimate association in their range ofoverlap in the Andes of Mendoza Province,Argentina. The hybrid resembles Calandriniacaespitosa more than Calandrinia compacta (seeTable I for comments regarding this hybrid). Theycf3-trnS sequence is that of the resident sample ofCalandrinia compacta, whereas the ITS isdimorphic. One ITS sequence corresponds toCalandrinia compacta and the other is possiblyrecombinatorial, with two positions shared by theresident Calandrinia caespitosa. The residentCalandrinia caespitosa shares the samedimorphism at the same two positions. (Thesedimorphisms were diagnosed by superimposedpeaks in the chromatograms. The ITS was notcloned.) The polymorphisms resolve as homoplasyin the Figure 4 topology. In the consensus of the100,000-plus ITS MP trees, all of the Calandriniacaespitosa and Calandrinia compacta branchescollapse into a polytomy. The lack of resolution isevident in the splits graph (Figure 7). Nonetheless,the combined data bootstrap supports both the


43

Calandrinia caespitosa and Calandrinia compactaclades, with the putative hybrid grouping with thelatter. It is also noteworthy that both the ITS andycf3-trnS sequences of a sample of Calandriniacompacta from Region III of Chile, north of the rangeof C. caespitosa, are different from those of the moresoutherly samples.

Finally, it is important to note that not all formsof Calandrinia have been sampled in the presentwork. Three recognized taxa not available areCalandrinia acaulis var. magna Macbride,Calandrinia alba (Ruiz & Pav.) DC., andCalandrinia acutisepala Añon. An internet searchreveals another species distributed worldwide byalpine garden enthusiasts as Calandriniaranunculina (nom. invalid.), originally from seedcollected by John M. Watson and Ana Flores inPatagonia. In addition, further study of Calandriniashould sample the entire ranges of especiallywidespread taxa and also intermediate formsassociated with two species pairs. One pair isCalandrinia acaulis and Calandrinia caroliniiHershk. & D.I.Ford. As noted by Hershkovitz & Ford(1993), many specimens from Peru and Bolivia aredifficult to assign to one or the other taxon. Theother pair is Calandrinia affinis Gillies ex Arn. andCalandrinia colchaguensis Barnéoud. These taxahave overlapping ranges in central Chile, and manyherbarium specimens are intermediates.

2. Calyptridium plus Cistanthe sectionsAmarantoides and Philippiamra. Because therelations of Cistanthe ambigua are unresolved, it isnot clear if the most recent common ancestor ofextant members of this clade was North American orSouth American, even though the sister group,Lenzia, is South American. Indeed, the relatively lowsequence divergence among samples of Cistanthesections Amarantoides and Philippiamra mayimplicate a secondary origin from North America (butsee also below).

Because of the low sequence divergence, thedata do not provide strong support for interspecificrelations among taxa of Cistanthe sectionsAmarantoides and Philippiamra, nor even thedistinction between these. This lack of divergenceis remarkable for a group traditionally circumscribedas including 14 species in two genera [12 spp. listedby Reiche (1898), including Cisanthe densiflora(Barnéoud) Carolin ex Hershk., plus Cistanthe

arancioana Peralta and Cistanthe minuscula (Añon)Peralta]. As discussed by Hershkovitz (1993), thesetaxa were at one time classified not only in separategenera, but in different tribes of the family. Thestrongest bootstrap value links Cistanthe calycinaof Cistanthe sect. Amarantoides with Cistantheamarantoides of Cistanthe sect. Philippiamra. Thedata do not even support monophyly of samples ofparticular species. The combined data provide at leastweak support for monophyly of the Chilean samplesof Cistanthe densiflora, but a specimen from SanJuan Province, Argentina, does not cluster with these.As noted in Table I, one collection appears to beintermediate between plants of Cistanthe calycinaand Cistanthe densiflora growing together in thesame locality. The possibility that gene flow canaccount for the apparent uniformity of sequences inthis and other genera is elaborated further below.

3. Cistanthe sect. Cistanthe3a. Grandiflora group. The taxonomy of the Grandi-flora group is difficult for reasons described in theMaterials and Methods, and because types of threespecies, including the type species of the genus, areiconotypes that illustrate plants cultivated in Europefrom seed of provenance no more specific than“Chile.” Given the tremendous variation amongpopulations, it may be necessary to designateneotypes. As indicated by the trees and splits graph,sequence divergence among the samples is low, butsomewhat conflictive. The graphics show proximityamong the samples of Cistanthe sp. indet., aff.Calandrinia laxiflora Phil., but the samplesthemselves were geographically proximal.

The combination of the data and field obser-vations suggest that interspecific gene flow may beoccurring in the Grandiflora group. For example,samples of Cistanthe sp. indet., aff. Calandrinialaxiflora and Cistanthe discolor (Schrad.) Spachcollected from the same site share both sequences.As here interpreted, the former species occurs strictlyon coastal cliffs and bluffs, has narrow and verysucculent leaves, petals with red blotches in the claw,and a well exserted style. The latter species rangesfrom coastal matorral to at least 50 km inland, hasbroad and flat leaves, whitish petal claws, and a stylenot exserted. I have observed intermediates wherethe two occur proximally, e.g., Cistanthe discolorwith petals that are red to maroon at the base andsomewhat exserted styles. One of the present

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samples of Cistanthe sp. indet., aff. Calandrinialaxiflora, Hershkovitz 99-948, has unusually broadleaves for this species, and plants in this populationhave supernumerary petals (6-8 rather than five).These anomalous characteristics may be indicativeof hybrid origin.

3b. Rosulatae group. As in the Grandiflora group,sequence divergence is low in Rosulatae group, andinterspecific relations generally are not resolved. Thepoor resolution of the sequence data is moresurprising for the Rosulatae group than for theGrandiflora group, because morphological andecological variation in the former group is greater.Reiche (1898) recognized 15 species divided amongthree sections of Calandrinia. Neither of the twosequences groups samples of some of the reaso-nably distinct species, let alone the morphologicallysimilar species corresponding to Reiche´s sections.The only exceptions are the partitions supportingrelationships of Cistanthe maritima, Cistanthecephalaphora, and Cistanthe sp. indet., aff.Calandrinia thyrsoidea. These last two taxa differfrom other members of the section in their congestedinflorescences of relatively small flowers with shortpedicels. They are nonetheless clearly distinct taxa.There does not appear to be a morphologicalcharacter that unites Cistanthe maritima with thesespecies, however. Cistanthe maritima, which isdisjunct in California, has the more openinflorescence and larger flowers of other membersof the section. However, support for monophyly ofthis partition depends upon the unresolved locationof the root, i.e., Cistanthe maritima may be the sisterof the remainder of the section.

As evident in Table I, most of the unidentifiedsamples fall into two morphological categories. Oneform resembles oversize plants of Cistanthearenaria. These commonly occur in association withother taxa, and I have found them only in highlydisturbed sites, especially in or near avocadoplantations. The other form somewhat resemblesCistanthe sp. indet., aff. Calandrinia thyrsoidea inhaving linear leaves and an erect rather than prostrateor ascending habit, but the flowers may be similar tothose of Cistanthe longiscapa or, alternatively,Cistanthe arenaria. These plants occur occasionallyin desert habitats in sites that may be disturbed butnot under cultivation.

As in the Grandiflora group, the molecular data,

morphology, and/or field observations suggestinterspecific gene flow in this group. The evidenceincludes observations of plants that are eitheraberrant or intermediate between other species, andthat occur in sympatry with potential parental typesand/or are restricted to human-transformed habitatsthat presumably were not part of the prehumanhistory of the group. The unidentified plantsdescribed above fit these criteria. As indicated inTable I, another aberrant collection, Cistanthe sp.02-80, resembles a dwarf or stunted plant of theGrandiflora group. The finding of both ITS and ycf3-trnS sequences of the Rosulatae group in thiscollection was somewhat surprising. In 2002, a fewhundreds of individuals were found near the peak ofCuesta Pajonales, along the main highway that linksthe cities of Coquimbo and Vallenar. The formapparently was first collected in the late 1950s, but ithas been collected perhaps three times since then.Cuesta Pajonales forms both a political and rain-shadow border between Regions III and IV of Chile.This is also a natural border of Chile’s “desiertoflorido”. In rainy years, vast expanses of Cistanthelongiscapa appear immediately on the Region IIIside of Cuesta Pajonales. This species is much moresparsely distributed on the Region IV side.Nonetheless, plants of both the Grandiflora and theRosulatae groups occur within 50 meters of thepopulation of aberrant plants. The aberrant plantsappeared to be fully fertile on the basis of theircopious seed production. Other aberrant plantsinclude those of giant-sized Cistanthe cymosa foundgrowing in a population of normal plants. The giantplants had ca. 17 stamens in contrast to the nearlyinvariable five that occur throughout the extensiverange of Cistanthe cymosa. Again, other forms ofthe Rosulatae group occur nearby. Another putativehybrid was an aberrant form of Cistanthe sp. indet.,aff. Calandrinia oblongifolia having pink rather thanusual white flowers and inordinately higher numbersof petals and stamens. The plants were growing withplants of Cistanthe sp. indet., aff. Calandriniaoblongifolia and Cistanthe humilis (Phil.) Peralta ina 100 m range of altitudinal overlap. Cistanthehumilis has pink flowers.

Molecular evidence for gene flow in theRosulatae group includes conflict among gene treesand conflict between gene trees and morphology,especially when such conflict is coincident withinterspecific sympatry. Conflict involving the gene


45

trees cannot be considered strong given the lowdivergence, i.e., it could be explained by homoplasyand/or conservation of ancestral polymorphism. Inany case, numerous conflicts exist between the ITSand ycf3-trnS trees for the Rosulatae group. Forexample, for the 12 samples of Cistanthe longiscapa,five can be considered to have conflicting combina-tions of genotypes, i.e., the different sequences donot circumscribe the same groups of samples. In onecase, a collection has the same ITS genotype of asympatric collection of Cistanthe cymosa. Thesequence of this Cistanthe cymosa sample is, in turn,different from the other four samples of this species.These collections are from a coastal site near thenorthern limits of the ranges of both species. Thissite is also near the type locality of Cistanthe cymosa.Another noteworthy example of discordance ispolyphyly of samples of Cistanthe arenaria in boththe ITS and ycf3-trnS sequences. Both trees showone sample grouping with a sample of Cistanthe sp.indet., aff. Calandrinia chamissoi, which Reiche(1898) considered a variety of Cistanthe arenaria.As identified here, the two taxa can be distinguishedby numerous characteristics (leaf shape, size of allfloral parts, sepal texture and coloration, petal color,stamen number, style length, stigma color, seedpubescence). Populations of each taxon occurseparately, but they commonly occur together,invariably with some morphologically intermediateindividuals (pers. obs.). Thus, the divergence of thematerial sampled here may reflect hybridization.

As a final note, the present analysis does notinclude 3-4 Peruvian taxa and one species fromGuadalupe Is., Mexico, all of which probably pertainto this section (see Macbride 1937; Hershkovitz1991a, 1991b; Hershkovitz & Zimmer 2000).

4. Montiopsis4a. Montiopsis subgenus Dianthoideae. This groupincludes at least four species. Two of these,Montiopsis andicola (Gillies) D.I.Ford andMontiopsis gayana, show considerable variability,as suggested by their extensive synonymy (Peralta1999). Likewise, each chloroplast haplotype ofsamples of these taxa was found to be distinct. Onespecies, Calandrinia tricolor, has been consideredas a synonym of Montiopsis andicola, but it isdistinct genetically, as well as morphologically(Hoffmann et al. 1998, p. 56, fig. 1 vs. fig. 4). It alsooccurs at lower elevation (pers. obs.) than

Montiopsis andicola. As in the other genera, theconflict between the ITS and ycf3-trnS sequencesmay be indicative of hybridization. However,additional sampling is needed to draw more definitiveconclusions.

4b. Montiopsis subgenus Montiopsis. As in othergenera, sequence divergence is too low to resolverelationships and evaluate conflicting signal. As inthe Rosulatae group of Cistanthe sect. Cistanthe,the lack of divergence is somewhat surprising giventhe morphological and ecological diversity of thetaxa, which Reiche (1897, 1898) also divided into threesections within Calandrinia. Ford (1992) recognized15 species. Notwithstanding, as in other genera,some evidence may suggest interspecific gene flow.The molecular data conflict in the relations of theMontiopsis trifida samples. The relevant partitionsreasonably well-supported in the ycf3-trnS tree areconflicted by the somewhat weaker partition in theITS tree. The partitions in the ycf3-trnS tree are lesswell supported in the combined data bootstrapconsensus. A more peculiar result is that onespecimen of Montiopsis parviflora (00-217) has thesame chloroplast haplotype as the Montiopsis trifidaspecimens and is well-separated from the remainingMontiopsis parviflora samples. This particularspecimen corresponds morphologically more to thetype specimen than the others. In particular, it hasdensely pubescent eglandular sepals rather thansparsely pubescent glandular sepals characteristicof the other samples. Also, its provenance is muchmore proximal to the type locality in northern RegionIV of Chile than the others, which are all from RegionIII. However, all of the samples of Montiopsisparviflora correspond morphologically and geogra-phically to the diagnosis given by Ford (1992).

The remaining evidence for gene flow inMontiopsis sect. Montiopsis is more circumstantial.A sample corresponding morphologically andgeographically to a putative recurring hybridbetween Montiopsis capitata (Hook. & Arn.)D.I.Ford and Montiopsis trifida (Ford 1992) has boththe ITS and ycf3-trnS sequences of the latter. Ford(1992) also suggested hybridization was responsiblefor intermediacy among the sympatric perennialspecies of Montiopsis, most of which are identicalin both their ITS and ycf3-trnS sequences. Thepresent work also documents a new case of sympatrybetween Montiopsis species, viz., Montiopsis

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modesta (Phil.) D.I.Ford (Hershkovitz 02-94) andMontiopsis parviflora (Hershkovitz 02-92, 02-93, &02-95). In fact, the Montiopsis parvifolia plants wereunusually diverse, and none of the three collectionshere have what Ford (1992) indicated was the“typical” morphology of the species. One collection(Hershkovitz 02-92) has a pedicel length longer thanthe extreme recorded by Ford and also plumose sepaltrichomes. The other two (Hershkovitz 02-93 andHershkovitz 02-95) have the sparely pubescent andglandular sepals that Ford regarded as unusual forthis species. This particular characteristic is sharedwith Montiopsis modesta, a species never beforecollected in the same elevational range nor floweringduring the same season as Montiopsis parviflora.

RAPID RADIATION, RAMPANT HYBRIDIZATION, OR BOTH?

A pervasive theme of the age of DNA sequencephylogenetics is that of “rapid radiation,” which canbe characterized as apparent morphological/ecologicaldiversity disproportionate to unexpectedly short andoften cladistically irresolvable gene tree branches. Theprevious analysis of western American PortulacaceaeITS data (Hershkovitz & Zimmer 2000) and aconsiderable number of references cited therein citedrapid radiation as the explanation for this pattern.Among the numerous more recent examples is Cronnet al. (2002). In most of these cases, rapid radiation isassociated with recency of the development of modernhabitats, especially on islands. The rapidity anddegree of morphological and ecological diversificationpossible in a short time and with negligible divergenceof “neutral marker” gene sequences is manifest byperforming BLAST searches of such domesticatedplants as Brassica species (pers. observ., data notshown).

Taken at face value, the present data can beinterpreted as evidence of successive rapid radiationsof taxa, the first resulting in evolution of the moderngenera and subsequent resulting in the speciesdiversity within each. At least with the sequencemarkers used, the radiation would be considered toorapid to evaluate the pattern of phylogeny across thevarious habitats occupied by the taxa. This isespecially the case for Montiopsis subg. Montiopsis,whose taxa exhibit the broadest range of habitats, fromcool temperate to desert and from high alpine to sealevel (Ford 1992). The present sequence data revealrelatively few differences among the taxa and practically

no phylogenetic resolution. A typical divergence ratefor ITS for herbaceous plants (5 x 10-9 substitutions/site/year; see Hershkovitz & Zimmer 2000) would datethe diversification of this group at less than 2 millionyears. This date is not inconceivable, but it suggeststhat the group diversified more recently than the range(though not necessarily the actual locations) ofhabitats occupied in Chile. In this case, the outgroupis clearly Montiopsis subg. Dianthoideae, whichincludes only alpine perennial species in central andsouthern Chile. This would suggest that Montiopsissubg. Montiopsis has such an origin and evolvednorthwards and into the lowland mediterranean, desert,and coastal regions. At the same time, it must be notedthat the ycf3-trnS divergence between the subgenerais relatively high, thus the rate calibration of one or theother or both sequences may be misleading. However,this does not change the observation that an order inwhich the species of Montiopsis subg. Montiopsisevolved into their diverse habitats cannot be discernedfrom either sequence, i.e., both sequences suggestsimul-taneous diversification. The same observationscan be extended to the other genera.

Possibly, permeation of the parsimony principle insystematics has promoted the rapid radiationinterpretation, viz., there is no need to postulate a DNAsequence diversification period longer than theminimum possible. The more recent implemen-tationof relatively simple likelihood methods inphylogenetics software, has facilitated a conceptualshift away from the “minimal explanation” towards theexplanation that best predicts the data under the mostrealistic model. In this context, it is possible to evaluate,at least intuitively and given certain parameters, analternative model. In particular, is it possible that theecological diversification within the genera ofPortulacaceae took place earlier than is apparent, butthat hybridization has homogeneized the neutralmarkers within genera, giving the appearance of morerecent radiation?

As a preface to consideration of the possible roleof hybridization (gene flow between historicallyindividuated lineages that nominally merit taxonomicdistinction) as an explanation for the morphologicaland molecular diversity of Andean Portulacaceae, itshould be emphasized that reticulate evolution in agroup predisposed to hybridization cannot beconsidered a priori an ad hoc assumption. In otherwords, it does not represent an explanation that isinherently arbitrary relative to other explanations. The


47

basis for an expectation of cladogenesis is reasonablyclear: the lack of panmixia in the face of continualmutation. In practice, however, cladogenesis de factorepresents a priori an ad hoc assumption, becausethe most popular phylogenetic methods constrain forcladogenesis without justification as to why thisprocess should be preferred, much less assumed, inthe group in question. It is perhaps more the rule thanthe exception that, at least in plants, the conditionsnecessary and sufficient for hybridization exist.Hybridization can be considered as a process that isnot necessary in organic evolution, but, like among-site rate heterogeneity, one that occurs at some level.In phylogenetics, hybridization is discriminated againstin part because it introduces tremendous mathematicalcomplexity into phylogenetic reconstruction and inpart because the relevant parameters are poorlyunderstood or cannot be easily evaluated usingcommon molecular systematic approaches (Linder &Rieseberg 2004). However, reticulate evolution andcladogenetic evolution can be evaluated subjectivelyin terms of several parameters, including the bioticconditions that regulate hybridization and whatgenotypic and phenotypic patterns are produced undereach model. Some of these parameters will be discussedbelow and evaluated in the case of AndeanPortulacaceae. In many cases, knowledge may not besufficient to establish or at least conjecture the valueof a particular parameter. The value of that parameter,however, should not be set at a value that, a priori,either favors or refutes hybridization or cladogenesis.

1. INTERFERTILITY

At present, no experimental data exist relating to theinterfertility among species within the genera ofPortulacaceae. However, several cases of suspectedhybrids sympatric with parental taxa have beendescribed above. It is also relevant to note that currentinterfertility may not be an indicator of past interfertility,i.e., fertility barriers are derived (Kimball et al. 2003).On the balance, the putative evidence for hybrids inthe absence of evidence for intersterility appears tofavor more than refute hybridization scenarios forPortulacaceae.

2. ALLOPOLYPLOIDY

Allopolyploidy is considered to be a consequence ofhybridization. However, hybrid taxa may also be diploid

(Rieseberg 1997). Among the relatively fewchromosome data for Portulacaceae (Nyanyano 1986;Ford 1992), there are no clear cases of allopolyploidy,but there are a few instances of tetraploids andhexaploids. Thus, the existing chromosome data shouldnot influence the hybridization probability one way oranother.

3. MORPHOLOGICAL EVIDENCE

Several cases of morphological intermediates havebeen discussed above. However, it should beemphasized that hybridization does not always resultin morphological intermediacy; hybrids may appearas one or the other parent or as a novel phenotype(McDade 1995, Rieseberg & Welch 2002). Likewise,some of the plants considered in the present analysisappeared to represent novelty.

4. CONFLICT BETWEEN GENOTYPES

The present results include several apparent cases ofconflict between ITS and ycf3-trnS genotypes, whichmay be interpreted as evidence of hybridization. Atthe same time, lack of conflict between two putativelyneutral markers cannot be considered strong evidencecontrary to hybridization. A large portion of the genomemay be neutral in terms of morphological appearance.Introgression of this portion across speciesboundaries is regulated mainly by interfertility/opportunity and perhaps fitness of the genotype atthe molecular level (Morjan & Rieseberg 2004,Rieseberg et al. 2004). Thus, while conflict betweenputatively neutral DNA sequences favors ahybridization scenario, agreement between two or afew markers may not yield strong evidence to thecontrary (see also Linder & Rieseberg 2004). Likewise,the paucity of ITS polymorphism in the present datamay not be contrary to hybrid origins. Elimination inhybrids of divergent parental ITS forms appears totake place over the course of relatively few generations,so that the inheritance is essentially uniparental inevolutionary time (Kotseruba et al. 2003, Song et al.1995, Smedmark et al. 2003, Kovarik, in press).

5. CONFLICT BETWEEN GENOTYPE AND MORPHOLOGY

Aside from cases in which suspected morphologicalhybrids lack polymorphism in their ITS sequence, twocases were noted in which samples have unexpected

Gayana Bot. 63(1), 2006

48

genotypes. One of these was a sample of Cistanthecymosa that shared the ITS genotype of a sympatricsample of Cistanthe longiscapa, and the other wassamples of Montiopsis parviflora with the ycf3-trnSsequence of Montiopsis trifida. The unexpectedsimilarity of the marker sequences in morphologicallydivergent taxa may itself result from hybridization.Introgression can cause alleles of one age to bereplaced by those of another. If younger alleles have atendency to replace older, the result would be lowsequence divergence in a group older than thesequence divergence would suggest. This can becalled marker “surfing,” because an allele can surf thetips of the phylogeny via introgression.

6. ECOLOGICAL CONDITIONS FAVORING HYBRIDS

If ecological instability and novelty of habitats favorhybrid success, then current and historicalenvironmental parameters must be consideredfavorable to hybridization of Andean Portulacaceae.Most of the diversity occurs in central and northernChile. This region includes altitudinal gradients ofcommonly 3000-4000 m over relatively short distances.The precipitation gradient ranges from absolute desertto moderate, and temperature regimes range fromsubtropical to boreal and maritime to continental. Thecombination of high relief and temperate latitudesprovide sharp climatic differences on opposing polarand equatorialfacing slopes. Chile has experiencedextreme ecological instability, especially in the periodof ca. 10 million – 10,000 ybp (Arroyo et al. 1988;Villagrán 1995; Villagrán & Hinojosa 1997; Hinojosa &Villagrán 2005). During this period, Chile´s extremealpine and desert habitats and its mediterranean typeclimate developed, these representing novelties forSouth America. The vegetation zones weresubstantially and repeatedly perturbed by advancingand retreating glaciers during the Pleistocene. Climaticinstability persists today, influenced by the SouthernOscillation, which generates cycles of significantclimate differences approximately every 5-10 years.These conditions collectively facilitate contactbetween species of different environments andpromote the availability of unoccupied habitats, bothof which favor hybridization.

7. SUMMARY

The above discussion is not intended to prove that

hybridization is the principal process underlyingdiversification of Portulacaceae in the Andeanregion. However, it does aim to show that frequenthybridization can yield the observed data patterns,that evidence for hybridization exists in some cases,and that evidence against hybridization is generallylacking. Thus, hypotheses of hybridization shouldnot be restricted to those instances where there isno other possible explanation. Evidence thathybridization may have been a recurring event duringthe evolution of Andean Portulacaceae suggests adifferent approach to taxonomic and phylogeneticstudies of these taxa. Floristic study must payattention to variability and aberrations in the field.Interpretation of molecular data derived from one ora few samples per taxon and few DNA sequencesmust allow for the possibility of past gene flow.Possibly more worthwhile would be approachesbased on multilocus techniques (e.g., AFLP) withsampling of multiple individuals from multiplepopulations. Empirical data on taxon interfertilitywould be useful. A program of artificial hybridizationmay to establish the extent to which phenotypicdiversity can be generated as a result of geneticsegregation (e.g., Carr et al. 1996, Schwarzbach etal. 2001). Such hybrids could be useful for compa-rison of their multilocus marker profiles withsuspected natural hybrids.

ACKNOWLEDGMENTS

I thank C. Zavaleta and C. Domínguez for technicaland/or field assistance, C. Hernández-Pellicer, A.Maldonado, and M. K. Arroyo (U. de Chile) and D.Hannon (RSA) for providing particular samples, R.Woods (Falklands Conservation Charity) forsamples and discussion, G. Arancio (ULS) forcollection information, the curators of CAS, RSA,SBBG, SD, UC, ULS for permission to sampleherbarium specimens, and Liz Zimmer and othermembers of the Laboratory of Systematic Biology,Smithsonian Institution and the staff of RanchoSanta Ana Botanical Garden for invaluable logisticalassistance in support of the development of amolecular systematics program in Chile. Themolecular research was undertaken in a laboratorydeveloped substantially with support from theMellon Foundation. The research was financed byCONICYT (Chile) grant number 1000909.


49

TAB

LE I.

Tax

a, sp

ecim

ens,

acro

nym

s, di

agno

stic

s, an

d co

mm

enta

ry fo

r pla

nt m

ater

ials

use

d in

this

stud

y. L

ocal

ity d

ata

prov

ided

is a

s com

plet

e as

is a

vaila

ble.

Dia

gnos

tics

and

com

men

tary

are

pro

vide

d pr

imar

ily in

the

case

whe

re p

ublis

hed

keys

and

des

crip

tions

are

insu

ffic

ient

and

/or i

n th

e ca

se o

f unu

sual

, pro

blem

atic

, or o

ther

wis

e no

tabl

eta

xa o

r spe

cim

ens.

Loca

lity

data

incl

ude

coun

try, f

irst p

oliti

cal d

ivis

ion,

and

usu

ally

seco

nd p

oliti

cal d

ivis

ion.

Exc

ept f

or th

e M

etro

polit

an R

egio

n, m

ain

polit

ical

div

isio

nsof

Chi

le ar

e ind

icat

ed w

ith R

oman

num

eral

s. Su

pers

crip

ts ad

jace

nt to

the a

cron

yms r

efer

to D

NA

extra

ctio

n pr

oced

ure:

1 ext

ract

ed fr

om h

erba

rium

mat

eria

l, 2 e

xtra

cted

from

silic

a-dr

ied

or fr

ozen

tiss

ue, 3 IT

S an

d/or

ycf3

-trnS

sequ

ence

s det

erm

ined

from

mor

e th

an o

ne in

depe

nden

t ext

ract

ion.

TAB

LA I.

La T

abla

pre

sent

a los

taxa

, esp

ecím

enes

, acr

ónim

os, d

iagn

óstic

os y

com

enta

rios d

e los

mat

eria

les u

sado

s en

este

estu

dio.

Los

dat

os d

e lo

calid

ad so

n en

trega

dos t

anco

mpl

etos

com

o fu

e pos

ible

. D

iagn

óstic

os y

com

enta

rios s

on en

trega

dos

prin

cipa

lmen

te en

el ca

so d

onde

las c

lave

s pub

licad

as y

las d

escr

ipci

ones

son

insu

ficie

ntes

y/o

enel

cas

o de

ser r

aras

o p

robl

emát

icas

, o p

ara

taxa

y e

spec

ímen

es n

otab

les.

Los d

atos

de

loca

lidad

incl

uyen

paí

s, re

gión

, pro

vinc

ia. E

xcep

to p

ara

la R

egió

n M

etro

polit

ana,

ladi

visi

ón p

olíti

ca p

rinci

pal d

e Chi

le se

indi

ca co

n nú

mer

os ro

man

os. E

l sup

erín

dice

junt

o a l

os ac

róni

mos

se re

fiere

n al

mét

odo

de ex

tracc

ión

de A

DN

: 1 ext

raíd

o de

sde m

ater

ial

de h

erba

rio, 2 e

xtra

ído

de te

jido

seca

do en

sílic

a o co

ngel

ado,

3 sec

uenc

ias d

e ITS

y/o

ycf3

-trnS

det

erm

inad

as d

e más

de u

na ex

tracc

ión

inde

pend

ient

e.

Gen

Ban

k se

quen

ceTa

xon

Acr

onym

Spec

imen

and

col

lect

ion

data

and

com

men

tary

acce

ssio

n(I

TS/y

cf3-

trnS

)

Cal

andr

inia

Kun

thC

alan

drin

ia se

ct. A

caul

es R

eich

eC

alan

drin

ia a

caul

is K

unth

acau

lis_1

3561

MEX

ICO

. Méx

ico:

Am

ecam

eca,

Gar

cía

1356

(RSA

)D

Q09

0306

/D

Q09

0285

acau

lis_2

7282

1M

EXIC

O. M

éxic

o: Z

inac

ante

pec,

Rze

dow

ski 2

7282

(RSA

)D

Q09

0307

/D

Q09

0286

Cala

ndrin

ia a

ffini

s Gill

ies e

x Arn

.af

finis

2,3

Vouc

her l

ost.

Orig

inal

dat

a: C

HIL

E. M

etro

polit

ana:

Cor

dille

ra, b

etw.

DQ

0903

18 /

Sant

iago

& V

alle

Nev

ado,

Her

shko

vitz

92-

05, d

eter

min

ed b

y M

. A.

DQ

0903

27H

ersh

kovi

tz.

Cal

andr

inia

car

olin

ii H

ersh

k. &

D. I

. For

dca

rolin

ii1A

RG

ENTI

NA

. Tuc

uman

: Que

brad

a lo

s B

emos

,D

Q09

0308

/Ar

royo

03-

108

(CO

NC

)D

Q09

0287

Cal

andr

inia

caes

pito

sa G

illie

s ex A

rn.

caes

pito

sa_7

902,

3C

ultiv

ated

. CH

ILE.

V:

ex F

ord

790

(MO

).D

Q09

0319

/D

Q09

0328

caes

pito

sa_2

6XI9

82,3

AR

GEN

TIN

A. M

endo

za: S

an C

arlo

s, rd

to L

agun

a D

iam

ante

,D

Q09

0305

/Pe

ralta

& H

ersh

kovi

tz s.n

. 26

Nov

199

2 (M

ERL)

DQ

0902

84

Gayana Bot. 63(1), 2006

50

Tabl

e I,

cont

inue

d

Gen

Ban

k se

quen

ceTa

xon

Acr

onym

Spec

imen

and

col

lect

ion

data

and

com

men

tary

acce

ssio

n(I

TS/y

cf3-

trnS

)

Cala

ndrin

ia ca

espi

tosa

x co

mpa

cta

x co

mpa

cta2,

3A

RG

ENTI

NA

. Men

doza

: San

Car

los,

rd to

Lag

una

Dia

man

te,

DQ

0903

09 /

Pera

lta &

Her

shko

vitz

s.n. 2

6 N

ov 1

992

(MER

L). C

OM

MEN

T:D

Q09

0288

The

pla

nt w

as s

ympa

tric

with

bot

h pu

tativ

e pa

rent

al s

peci

es. I

t had

am

assi

ve c

row

n of

thic

kene

d st

ems r

esem

blin

g th

at o

f C. c

aesp

itosa

rath

er th

en th

e sl

ende

r rhi

zom

es o

f C. c

ompa

cta.

The

pet

als a

re ro

sera

ther

than

ora

nge

as in

C. c

aesp

itosa

or p

ale

pink

as i

n C

. com

pact

a.Th

e le

aves

and

ped

uncl

es a

re in

term

edia

te in

leng

th b

etw

een

the

two

pare

ntal

spe

cies

.

Cala

ndrin

ia co

lcha

guen

sis B

arné

oud

colc

hagu

ensi

s2,3

CH

ILE.

VII

I: Ñ

uble

, Ter

mas

de

Chi

llán,

dry

rock

y sl

opes

E o

fD

Q09

0303

/ho

tel/s

pa, H

ersh

kovi

tz 0

1-72

(C

ON

C).

DQ

0902

82

Cal

andr

inia

com

pact

a B

arné

oud

com

pact

a_01

-512

CH

ILE.

III:

Hua

sco,

rd fr

om C

onay

to E

l Nev

ado

(Bar

rick

Min

e Pr

oyec

toD

Q09

0310

/Pa

scua

) at k

m 3

6, H

ersh

kovi

tz 0

1-51

(CO

NC

).D

Q09

0289

com

pact

a_26

XI9

82,3

AR

GEN

TIN

A. M

endo

za: S

an C

arlo

s, rd

to L

agun

a D

iam

ante

,D

Q09

0262

/Pe

ralta

& H

ersh

kovi

tz s.n

. 26

Nov

199

2 (M

ERL)

DQ

0903

29

Cal

andr

inia

sp.

inde

t. 1

C. s

p_98

-22B

2C

HIL

E. M

etro

polit

ana:

Cor

dille

ra, V

alle

Nev

ado

ski a

rea,

DQ

0903

04 /

Her

shko

vitz

98-

22B

(CO

NC

).D

Q09

0283

Cal

andr

inia

sect

. Cal

andr

inia

Cala

ndrin

ia a

xilli

flora

Bar

néou

dax

illifl

ora_

99-3

7741

CH

ILE.

Arr

oyo

99-3

774

(CO

NC

)D

Q09

0290

/D

Q09

0266

axill

iflor

a_99

-385

21C

HIL

E. A

rroy

o 99

-385

2 (C

ON

C)

DQ

0902

91 /

DQ

0902

67

Cal

andr

inia

bre

wer

i S. W

atso

nbr

ewer

i_26

21U

SA. C

alifo

rnia

: San

Die

go C

o., S

Cuy

amac

a M

ts, H

irsc

hber

g 26

2 (R

SA)

DQ

0902

96 /

DQ

0902

71


51

Tabl

e I,

cont

inue

dG

enB

ank

sequ

ence

Taxo

nA

cron

ymSp

ecim

en a

nd c

olle

ctio

n da

ta a

nd c

omm

enta

ryac

cess

ion

(ITS

/ycf

3-tr

nS)

brew

eri_

2501

1U

SA. C

alifo

rnia

: Sta

. Cru

z I.,

Jun

ak S

C 2

501

(SB

BG

)D

Q09

0295

/D

Q09

0272

brew

eri_

4204

1U

SA. C

alifo

rnia

: San

Lui

s O

bisp

o C

o., S

anta

Luc

ia M

ts, J

unak

420

4 (R

SA)

DQ

0902

65 /

DQ

0902

73

brew

eri_

4289

1,3

USA

. Cal

iforn

ia: S

an L

uis O

bisb

o C

o., J

unak

428

9 (S

BB

G)

DQ

0902

63 /

DQ

0902

74

brew

eri_

8425

1U

SA. C

alifo

rnia

: Riv

ersi

de C

o., P

alom

ar R

ange

, Agu

a Tib

ia W

ild. A

rea,

DQ

0902

64 /

Boyd

842

5 (R

SA)

DQ

0902

75

Cal

andr

inia

cilia

ta (R

uiz &

Pav

.) D

Cci

liata

_031

071

AR

GEN

TIN

A. T

úcum

an: Q

uebr

ada

los

Bem

os, A

rroy

o 03

-107

(CO

NC

)D

Q09

0292

/D

Q09

0268

cilia

ta_1

5501

MEX

ICO

. Tla

xcal

a: N

anac

amilp

a, R

odrí

guez

155

0 (R

SA)

DQ

0902

93 /

DQ

0902

69

cilia

ta_4

0430

1,3

MEX

ICO

. Chi

apas

: Mot

ozin

tla d

e M

endo

za, 3

000

m,

DQ

0902

94 /

Bree

dlov

e 404

30 (R

SA)

DQ

0902

70

Cal

andr

inia

com

pres

sa S

chra

d. ex

DC

com

pres

sa2,

3Vo

uche

r los

t. O

rigin

al d

ata:

CH

ILE.

VII

I: B

ío-B

ío, r

d be

tw. C

once

pció

n &

DQ

0903

14 /

Yum

bel c

a. 1

km

W o

f jct

w/rd

to C

abre

ro, H

ersh

kovi

tz 9

2-10

,D

Q09

0323

dete

rmin

ed b

y M

. A. H

ersh

kovi

tz

Cal

andr

inia

men

zies

ii (H

ook.

)m

enzi

esii_

02-2

02U

SA. C

alifo

rnia

: Am

ador

Co.

, Cal

iforn

ia H

wy

88 a

t jct

w/Ir

ish

DQ

0902

97 /

Torr.

& A

. Gra

yTo

wn

Rd

& C

linto

n R

d, H

ersh

kovi

tz 0

2-20

DQ

0902

76

men

zies

ii_24

9991

CA

NA

DA

. Bri

tish

Col

umbi

a: V

anco

uver

I., C

alde

r 29

499

(RSA

)D

Q09

0298

/D

Q09

0281

Gayana Bot. 63(1), 2006

52

Tabl

e I,

cont

inue

d

Gen

Ban

k se

quen

ceTa

xon

Acr

onym

Spec

imen

and

col

lect

ion

data

and

com

men

tary

acce

ssio

n(I

TS/y

cf3-

trnS

)

men

zies

ii_25

791

MEX

ICO

. Baj

a C

alifo

rnia

Nor

te: R

ebm

an 2

579

(RSA

)D

Q09

0299

/D

Q09

0277

men

zies

ii_25

5601

USA

. Ari

zona

: Pim

a C

o, H

itchc

ock

2556

0 (R

SA)

DQ

0903

00 /

DQ

0902

78

men

zies

ii_25

921

USA

. Ari

zona

: Pim

a C

o., C

oron

ado

Nat

iona

l For

est,

Schm

idt 2

592

(RSA

)D

Q09

0301

/D

Q09

0279

men

zies

ii_44

591

USA

. Ore

gon:

Lin

n C

o., H

alse

445

9 (R

SA)

DQ

0903

02 /

DQ

0902

80

men

zies

ii_IV

1992

2,3

No

vouc

her.

Orig

inal

dat

a: U

SA. C

alifo

rnia

: Yub

a C

o.,

DQ

0903

15 /

Her

shko

vitz

s.n

. Apr

199

2, d

eter

min

ed b

y M

. A. H

ersh

kovi

tzD

Q09

0324

men

zies

ii_M

al11

No

vouc

her.

Orig

inal

dat

a: C

ultiv

ated

, Uni

ted

Kin

gdom

. Fal

klan

d Is

land

s:D

Q09

0316

/St

anle

y. F

ragm

ents

of t

he p

lant

wer

e re

ceiv

ed fr

om D

r. D

avid

Bro

ught

on,

DQ

0903

25R

oyal

Bot

anic

Gar

den,

Kew

men

zies

ii_M

al21

No

vouc

her,

Orig

inal

dat

a: C

ultiv

ated

, Uni

ted

Kin

gdom

.D

Q09

0317

/Fa

lkla

nd Is

land

s: S

tanl

ey, e

x se

ed fr

om B

eave

r Isl

and,

S. P

once

t,D

Q09

0326

s.n. F

ragm

ents

of t

he p

lant

wer

e re

ceiv

ed fr

om D

r. D

avid

Bro

ught

on,

Roy

al B

otan

ic G

arde

n, K

ew

Cal

andr

inia

sp. i

ndet

. 2C

_sp_

Mal

11N

o vo

uche

r, O

rigin

al d

ata:

Cul

tivat

ed, U

nite

d K

ingd

om. F

alkl

and

Isla

nds:

DQ

0903

12 /

Stan

ley,

ex

seed

from

Tea

Isla

nd, S

. Pon

cet,

s.n. F

ragm

ents

of t

he p

lant

DQ

0903

21w

ere

rece

ived

from

Dr.

Dav

id B

roug

hton

, Roy

al B

otan

ic G

arde

n, K

ew

C_s

p_M

al21

No

vouc

her.

Orig

inal

dat

a: U

nite

d K

ingd

om. F

alkl

and

Isla

nds.:

NE

faci

ngD

Q09

0313

/sl

ope

of S

abin

a Po

int,

at h

ead

of N

orth

Har

bor,

R. W

oods

s.n.

Dec

. 199

9 .

DQ

0903

22D

NA

ext

ract

ed fr

om se

ed se

nt b

y R

. Woo

ds, F

alkl

ands

Con

serv

atio

n C

harit

y.


53

Tabl

e I,

cont

inue

d

Gen

Ban

k se

quen

ceTa

xon

Acr

onym

Spec

imen

and

col

lect

ion

data

and

com

men

tary

acce

ssio

n(I

TS/y

cf3-

trnS

)

Ca

land

rinia

sect

. Mon

ocos

mia

(Fen

zl)

H

ersh

k.C

alan

drin

ia m

onan

dra

DC

mon

andr

a2Vo

uche

r los

t. O

rigin

al d

ata:

CH

ILE.

VII

I: Ñ

uble

, Fun

do L

as S

alvi

das,

DQ

0903

11 /

Her

shko

vitz

92-

08, d

eter

min

ed b

y M

. A. H

ersh

kovi

tzD

Q09

0320

Cal

yptr

idiu

m N

utt.

Cal

yptr

idiu

m m

onan

drum

Nut

t.m

onan

drum

2U

SA. C

alifo

rnia

: San

Ber

nard

ino

Co.

, Cal

iforn

ia H

wy

138

ca. 1

km

DQ

0903

72 /

W o

f US

Inte

rsta

te H

wy

15, H

ersh

kovi

tz 0

2-01

(CO

NC

).D

Q09

0338

Cal

yptr

idiu

m m

onos

perm

um G

reen

em

onos

perm

um2

Vouc

her l

ost.

Orig

inal

dat

a: U

SA. C

alifo

rnia

: Alp

ine

Co.

, alo

ng H

wy

DQ

0903

75 /

88 S

ierr

a N

evad

a, H

ersh

kovi

tz s.

n. A

pr. 1

992,

det

erm

ined

by

M. A

.D

Q09

0341

Her

shko

vitz

Cal

yptr

idiu

m m

onos

perm

um x

um

bella

tum

x m

onos

perm

um2

USA

. Ore

gon:

Jack

son

Co.

, Mt.

Ash

land

Rd

ca. 0

.5 m

iles E

of s

ki lo

dge,

DQ

0903

67 /

Her

shko

vitz

02-

38 (

CO

NC

).D

Q09

0333

Cal

yptr

idiu

m p

arry

i A. G

ray

var.

neva

dens

epa

rryi

nev

aden

se1

USA

. Nev

ada:

Nye

Co.

, Tie

hm 1

3605

(CA

S)D

Q09

0374

/J.

T. H

owel

lD

Q09

0340

Cal

yptr

idiu

m p

arry

i var

. par

ryi

parr

yi p

arry

i2U

SA. C

alifo

rnia

: San

Ber

nard

ino

Co.

, San

Ber

nard

ino

Nat

l. Fo

rest

,D

Q09

0371

/H

ersh

kovi

tz 0

2-11

(C

ON

C).

DQ

0903

37

Cal

yptr

idiu

m p

arry

i var

. ari

zoni

cum

parr

yiM

EXIC

O. B

aja

Cal

iforn

ia N

orte

: Reb

man

417

4 (S

D)

DQ

0903

73 /

J. T.

How

ell

ariz

onic

um_4

1741

DQ

0903

39

parr

yiVo

uche

r los

t. O

rigin

al d

ata:

MEX

ICO

. Son

ora:

Cer

ro C

olor

ado,

DQ

0903

71 /

ariz

onic

um_2

2III

931

Piña

rte R

egió

n, F

elge

r s.n

. 22

Mar

199

3D

Q09

0330

Cal

yptr

idiu

m p

ulch

ellu

m (E

astw

.) H

oove

rpu

lche

llum

1U

SA. C

alifo

rnia

: Mad

era

Co.

, Ham

on 8

0-67

(UC

)D

Q09

0370

/D

Q09

0336

Gayana Bot. 63(1), 2006

54

Tabl

e I,

cont

inue

d

Gen

Ban

k se

quen

ceTa

xon

Acr

onym

Spec

imen

and

col

lect

ion

data

and

com

men

tary

acce

ssio

n(I

TS/y

cf3-

trnS

)

Cal

yptr

idiu

m p

ygm

aeum

Par

ish

ex R

ydb.

pygm

aeum

1U

SA. C

alifo

rnia

: Tw

isse

lman

168

11 (R

SA)

DQ

0903

68 /

DQ

0903

34

Cal

yptr

idiu

m q

uadr

ipet

alum

S. W

atso

nqu

adrip

etal

um2,

3Vo

uche

r los

t. O

rigin

al d

ata:

USA

. Cal

iforn

ia: L

ake

Co.

, Her

shko

vitz

DQ

0903

65 /

s.n. A

pr. 1

992,

det

erm

ined

by

M. A

. Her

shko

vitz

DQ

0903

31

Cal

yptr

idiu

m ro

seum

S. W

atso

nro

seum

USA

. Nev

ada:

Lyo

n C

o., H

ersh

kovi

tz 0

2-25

(CO

NC

).D

Q09

0366

/D

Q09

0332

Cal

yptr

idiu

m u

mbe

llatu

m (T

orr.)

Gre

ene

umbe

llatu

m1

USA

. Cal

iforn

ia: M

ono

Co.

, Big

San

d Fl

at, H

oner

132

(RSA

)D

Q09

0369

/D

Q09

0335

Cista

nthe

Spa

ch

C

ista

nthe

sect

. Am

aran

toid

es

R

eich

e (H

ersh

k.)

Cis

tant

he a

mbi

gua

(S. W

atso

n)am

bigu

a1U

SA. C

alifo

rnia

: San

Ber

nard

ino

Co.

, Han

non

& E

lvin

s.n.

(RSA

)D

Q09

0396

/C

arol

in ex

Her

shk.

DQ

0903

62

Cista

nthe

caly

cina

(Phi

l.) C

arol

in ex

Her

shk.

caly

cina

_00-

1702

CH

ILE.

III:

Hua

sco,

rd fr

om V

alle

nar t

o Ju

nta V

aler

iana

40

km fr

omD

Q09

0387

/ct

r of V

alle

nar,

Her

shko

vitz

00-

170

(CO

NC

).D

Q09

0353

caly

cina

_00-

1942

CH

ILE.

III:

Hua

sco,

rd fr

om D

omey

ko to

Cal

eta

Sarc

o 2

km W

DQ

0903

82 /

of D

omey

ko, H

ersh

kovi

tz 0

0-19

4 (C

ON

C).

DQ

0903

47

caly

cina

_00-

202

CH

ILE.

II: T

alta

l, km

114

0 of

Pan

amer

ican

Hw

y, b

etw.

Ant

ofag

asta

DQ

0903

82 /

& T

alta

l, H

ersh

kovi

tz 0

0-20

(CO

NC

).D

Q09

0348

Cist

anth

e sal

solo

ides

(Bar

néou

d)sa

lsol

oide

s2C

HIL

E. II

: Ant

ofag

asta

, rd

from

Ant

ofag

asta

to M

ina

Esco

ndid

a be

tw. k

mC

arol

in ex

Her

shk.

131

& 1

35, H

ersh

kovi

tz 0

0-04

(CO

NC

).

Cis

tant

he sp

. ind

et. 1

, aff

. C. c

alyc

ina

aff.

caly

cina

2C

HIL

E. IV

: Elq

ui, R

uta

41 c

a. 3

6 km

E o

f Vic

uña,

Her

shko

vitz

00-

227

DQ

0903

78 /


55

Tabl

e I,

cont

inue

dG

enB

ank

sequ

ence

Taxo

nA

cron

ymSp

ecim

en a

nd c

olle

ctio

n da

ta a

nd c

omm

enta

ryac

cess

ion

(ITS

/ycf

3-tr

nS)

(CO

NC

). C

OM

MEN

T: T

he h

abit

of th

is p

lant

was

inte

rmed

iate

bet

wee

n th

eD

Q09

0344

pros

trate

form

of C

. cal

ycin

a an

d th

e as

cend

ing

form

C. d

ensi

flora

, bot

h of

whi

ch g

row

at t

his s

ite.

Cista

nthe

den

siflo

ra (B

arné

oud)

Car

olin

dens

iflor

a_00

-101

2C

HIL

E. II

I: H

uasc

o, P

anam

eric

an H

wy

betw

. La S

eren

a & V

alle

nar c

a.D

Q09

0386

/ex

Her

shk.

69 k

m fr

om V

alle

nar,

Her

shko

vitz

00-

101

(CO

NC

).D

Q09

0352

dens

iflor

a_00

-171

2C

HIL

E. II

I: H

uasc

o, rd

from

Dom

eyko

to C

alet

a Sa

rco

2 km

W o

f Dom

eyko

,D

Q09

0383

/H

ersh

kovi

tz 0

0-17

1 (C

ON

C).

DQ

0903

49

dens

iflor

a_00

-226

2C

HIL

E. IV

: Elq

ui, R

uta

41 c

a. 3

6 km

E o

f Vic

uña,

Her

shko

vitz

00-

226

DQ

0903

84 /

DQ

0903

50

dens

iflor

a_00

-952

CH

ILE.

III:

Cop

iapó

, rd

from

Pai

pote

to D

iego

de A

lmag

ro c

a. 9

1 km

ND

Q09

0385

/of

Pai

pote

, Her

shko

vitz

00-

95 (C

ON

C).

DQ

0903

51

Cis

tant

he sp

. ind

et. 2

, aff

. C. d

ensi

flora

aff.

dens

iflor

a2,3

AR

GEN

TIN

A. S

an J

uan:

ex

Kie

sling

& P

eral

ta s.

n. (

MER

L)D

Q09

0376

/D

Q09

0342

Cis

tant

he se

ct. C

ista

nthe

CO

MM

ENT:

Whe

n no

ted,

see

d su

rfac

e m

orph

olog

y is

as

follo

ws:

G,

glab

rous

; H, h

airy

; PT,

pus

ticul

ate-

tom

ento

se.

Gra

ndifl

ora

grou

p (=

Cal

andr

inia

sect

.Ci

stant

he R

eich

e)C

ista

nthe

cab

rera

e (A

ñon)

I. P

eral

taca

brer

ae2

CH

ILE.

III:

Hua

sco,

rd fr

om C

onay

to E

l Nev

ado

(Bar

rick

Min

eD

Q09

0250

/Pr

oyec

to P

ascu

a) c

a. 5

km

S o

f Con

ay, H

ersh

kovi

tz 0

1-33

(CO

NC

).D

Q09

0185

Cista

nthe

disc

olor

(Sch

rad.

) Spa

chdi

scol

or_0

3-12

2C

HIL

E. V

: Aco

ncag

ua, E

of C

abild

o, H

ersh

kovi

tz 0

3-12

(CO

NC

).D

Q09

0261

/D

Q09

0195

disc

olor

_03-

282

CH

ILE.

V: A

conc

agua

, rd

betw

. Pap

udo

& Z

apal

lar,

Her

shko

vitz

DQ

0902

57 /

03-2

8 (C

ON

C).

DQ

0901

91

Gayana Bot. 63(1), 2006

56

Tabl

e I,

cont

inue

d

Gen

Ban

k se

quen

ceTa

xon

Acr

onym

Spec

imen

and

col

lect

ion

data

and

com

men

tary

acce

ssio

n(I

TS/y

cf3-

trnS

)

Cis

tant

he g

rand

iflor

a (L

indl

ey)

gran

diflo

ra_9

9-94

52C

HIL

E. IV

: Cho

apa,

Pan

amer

ican

Hw

y ca

. 3 k

m N

of L

os V

ilos,

DQ

0902

46 /

Car

olin

ex H

ersh

k.H

ersh

kovi

tz 9

9-94

5 (C

ON

C).

CO

MM

ENT:

See

ds H

.D

Q09

0181

gran

diflo

ra_9

9-97

72C

HIL

E. M

etro

polit

ana:

Cha

cabu

co, r

d fr

om T

iltil

to L

imac

he c

a. 2

km

ED

Q09

0240

/of

Pue

nte

Sant

a La

ura,

Her

shko

vitz

99-

977

(CO

NC

). C

OM

MEN

T: S

eeds

H.

DQ

0901

75

Cis

tant

he sp

. ind

et. 3

, aff

.cr

assi

folia

2C

HIL

E. II

I: H

uasc

o, je

ep tr

ail c

a. 3

0 ai

r km

N o

f Car

rizal

Baj

o, H

ersh

kovi

tzD

Q09

0247

/C

alan

drin

ia cr

assi

folia

Phi

l.00

-123

(CO

NC

).D

Q09

0182

Cis

tant

he sp

. ind

et. 4

, aff

.la

xiflo

ra_9

9-94

82C

HIL

E. IV

: Cho

apa,

Pan

amer

ican

Hw

y ca

. 5 k

m N

of L

os V

ilos,

Her

shko

vitz

DQ

0902

45 /

Cala

ndrin

ia la

xiflo

ra P

hil.

99-9

48 (C

ON

C).

CO

MM

ENT:

See

ds H

.D

Q09

0180

laxi

flora

_03-

142

CH

ILE.

IV: C

hoap

a, P

anam

eric

an H

wy

ca. 5

km

N o

f Los

Vilo

s, H

ersh

kovi

tzD

Q09

0254

/03

-14

(CO

NC

).D

Q09

0189

laxi

flora

_03-

232

CH

ILE.

V: A

conc

agua

, Pan

amer

ican

Hw

y 2

km N

of H

uenq

uen,

Her

shko

vitz

DQ

0902

55 /

03-2

3 (C

ON

C).

DQ

0901

90

laxi

flora

_03-

292

CH

ILE.

V: A

conc

agua

, rd

betw

. Pap

udo

& Z

apal

lar,

2km

N o

f N a

cces

s to

DQ

0902

58 /

Zapa

llar,

Her

shko

vitz

03-

29 (C

ON

C).

DQ

0901

92

laxi

flora

_03-

302

CH

ILE.

V: A

conc

agua

, rd

betw

. Pap

udo

& Z

apal

lar,

2km

N o

f N a

cces

s to

DQ

0902

59 /

Zapa

llar,

Her

shko

vitz

03-

30 (C

ON

C).

DQ

0901

93

laxi

flora

_03-

372

CH

ILE.

V: A

conc

agua

, Hor

cón,

Her

shko

vitz

03-

37 (C

ON

C).

DQ

0902

60 /

DQ

0901

94

Cis

tant

he sp

. ind

et. 5

, aff

. Cal

andr

inia

muc

ronu

lata

2C

HIL

E. V

II: C

auqu

enes

, Con

stitu

ción

, Her

shko

vitz

03-

194

(CO

NC

).D

Q09

0251

/m

ucro

nula

ta M

eyen

DQ

0901

86


57

Tabl

e I,

cont

inue

d

Gen

Ban

k se

quen

ceTa

xon

Acr

onym

Spec

imen

and

col

lect

ion

data

and

com

men

tary

acce

ssio

n(I

TS/y

cf3-

trnS

)

Cis

tant

he sp

. ind

et. 6

, aff

. Cal

andr

inia

spec

iosa

2C

HIL

E. II

I: C

opia

pó, r

d fr

om P

aipo

te to

Die

go d

e Alm

agro

ca.

9 k

m N

DQ

0902

43 /

spec

iosa

Leh

m.

of P

aipo

te, H

ersh

kovi

tz 0

0-91

(CO

NC

).D

Q09

0178

Cis

tant

he sp

. ind

et. 7

C. s

p_00

-114

2C

HIL

E. II

I: C

opia

pó, P

anam

eric

an H

wy

betw

. Cop

iapó

& V

alle

nar c

a.D

Q09

0241

/40

km

S o

f Cop

iapó

, Her

shko

vitz

00-

114

(CO

NC

). C

OM

MEN

T: s

eeds

H.

DQ

0901

76

C. s

p_00

-214

2C

HIL

E. IV

: Elq

ui, r

d be

twe.

Mar

ques

a &

Con

doria

co k

m 1

2 N

of M

arqu

esa,

DQ

0902

44 /

Her

shko

vitz

00-

214

(CO

NC

). C

OM

MEN

T: s

eeds

H.

DQ

0901

79

C. s

p_00

-312

CH

ILE.

II: T

alta

l, rd

bet

w. T

alta

l & P

apos

o ca

. 20

km N

of T

alta

l, H

ersh

kovi

tzD

Q09

0249

/00

-31

(CO

NC

). C

OM

MEN

T: s

eeds

H.

DQ

0901

84

C. s

p_00

-512

CH

ILE.

II: T

alta

l, rd

bet

w. T

alta

l & P

apos

o ca

. 26

km N

of T

alta

l, H

ersh

kovi

tzD

Q09

0248

/00

-51.

CO

MM

ENT:

sta

men

s ca

. 100

, see

ds G

, col

lect

ed w

ith H

ersh

kovi

tzD

Q09

0183

00-6

1 (C

ON

C).

C. s

p_00

-612

CH

ILE.

II: T

alta

l, rd

bet

w. T

alta

l & P

apos

o ca

. 26

km N

of T

alta

l, H

ersh

kovi

tzD

Q09

0242

/00

-61

(CO

NC

). C

OM

MEN

T: s

tam

ens

ca. 5

0, s

eeds

H, c

olle

cted

with

DQ

0901

77H

ersh

kovi

tz 0

0-51

.

C. s

p_00

-144

2C

HIL

E. IV

: Elq

ui, P

anam

eric

an H

wy

betw

. Los

Vilo

s & L

a Ser

ena c

a. 1

km

ND

Q09

0253

/of

Las

Tac

as jc

t, H

ersh

kovi

tz 0

0-14

4 (C

ON

C).

CO

MM

ENT:

Pla

nts

DQ

0901

88re

sem

ble C

ista

nthe

gra

ndifl

ora

exce

pt fo

r the

ir un

usua

lly b

road

and

lax

leav

es.

Ros

ulat

ae g

roup

And

inae

form

s (=

Cal

andr

inia

sect

.

An

dina

e R

eich

e)C

ista

nthe

frig

ida

(Bar

néou

d) I.

Per

alta

frigi

da2

CH

ILE.

Met

ropo

litan

a: C

ordi

llera

, Val

le N

evad

o sk

i are

a, H

ersh

kovi

tzD

Q09

0215

/01

-28

(CO

NC

).D

Q09

0150

Gayana Bot. 63(1), 2006

58

Tabl

e I,

cont

inue

d

Gen

Ban

k se

quen

ceTa

xon

Acr

onym

Spec

imen

and

col

lect

ion

data

and

com

men

tary

acce

ssio

n(I

TS/y

cf3-

trnS

)

Cis

tant

he h

umili

s (Ph

il.) I

. Per

alta

hum

ilis2

CH

ILE.

IV: E

lqui

, Cor

dille

ra D

oña A

na, F

undo

Oliv

iere

s, H

ersh

kovi

tz 0

1-17

DQ

0901

98(C

ON

C).

CO

MM

ENT:

Flo

wer

s 10

-15

per i

nflo

resc

ence

, sta

men

s 20

-25,

DQ

0901

33se

eds P

T. T

he id

entif

icat

ion

of th

is p

lant

is b

ased

mai

nly

on th

e or

igin

of t

hety

pe ([

F. P

eral

ta s.

n. (S

GO

!)] “

in a

ltiss

imo

mon

te d

oña A

na, c

. 400

0 m

.s.m

.)an

d on

supe

rfic

ial s

imila

rity

to th

e ty

pe v

eget

ativ

ely.

For

d-W

ernt

z &

Per

alta

(200

2) in

dica

ted

few

er fl

ower

s (5-

8), f

ewer

stam

ens (

5-6)

, and

gla

brou

s see

dsfo

r thi

s sp

ecie

s.

Cis

tant

he sp

. ind

et. 8

, aff

. Cal

andr

inia

oblo

ngifo

lia2

CH

ILE.

IV: E

lqui

, Cor

dille

ra D

oña A

na, F

undo

Oliv

iere

s, H

ersh

kovi

tzD

Q09

0197

/ob

long

ifolia

Bar

néou

d 0

1-15

(CO

NC

).D

Q09

0132

Cista

nthe

hum

ilis x

Cal

andr

inin

iax

oblo

ngifo

lia2

CH

ILE.

IV: E

lqui

, Cor

dille

ra D

oña A

na, F

undo

Oliv

iere

s, H

ersh

kovi

tzD

Q09

0196

/ob

long

ifolia

01-1

6 (C

ON

C).

CO

MM

ENT:

The

se p

lant

s oc

cur i

n th

e el

evat

iona

l zon

eD

Q09

0131

of o

verla

p be

twee

n th

e pu

tativ

e pa

rent

al ta

xa. T

hey

supe

rfic

ially

rese

mbl

eC

alan

drin

ia o

blon

gifo

lia, b

ut h

ave

pink

pet

als o

f the

Cis

tant

he h

umili

s rat

her

than

whi

te o

f the

form

er, a

s wel

l as s

ever

al fl

oral

cha

ract

eris

tics d

istin

ct fr

om b

oth

puta

tive

pare

ntal

form

s.

Cis

tant

he sp

. ind

et. 9

, aff

. Cal

andr

inia

villa

nuev

ae1

No

vouc

her.

Orig

inal

dat

a: C

HIL

E. II

-III

: Rd

betw

. Ex

Ofic

ina F

lor d

e Chi

leD

Q09

0199

/vi

llanu

evae

Phi

l. &

Sal

ar d

e la

Azu

frer

a, A

. Mal

dona

do s.

n., d

eter

min

ed b

y M

. A. H

ersh

kovi

tzD

Q09

0134

Are

narie

gro

up (=

Cal

andr

inia

sect

.

A

rena

rie

Rei

che)

Cis

tant

he a

rena

ria

(Cha

m.)

Car

olin

aren

aria

_00-

2152

CH

ILE.

IV: E

lqui

, rd

betw

. Mar

ques

a &

Con

doria

co k

m 1

2 N

of M

arqu

esa,

DQ

0902

11 /

ex H

ersh

k.H

ersh

kovi

tz 0

0-21

5 (C

ON

C).

DQ

0901

46

aren

aria

_00-

2512

CH

ILE.

IV: E

lqui

, rd

from

Vic

uña

to H

urta

do 1

0.5

km S

of j

ct w

/Rut

a 41

,D

Q09

0233

/H

ersh

kovi

tz 0

0-25

1 (C

ON

C).

DQ

0901

74

Cis

tant

he sp

. ind

et 1

0, a

ff. C

alan

drin

iach

amis

soi2

CH

ILE.

IV: C

hoap

a, rd

bet

w. Il

lape

l & C

omba

rbal

a, H

ersh

kovi

tz .

DQ

0902

39 /

cham

isso

i Bar

néou

d99

-870

(CO

NC

)D

Q09

0173


59

Tabl

e I,

cont

inue

dG

enB

ank

sequ

ence

Taxo

nA

cron

ymSp

ecim

en a

nd c

olle

ctio

n da

ta a

nd c

omm

enta

ryac

cess

ion

(ITS

/ycf

3-tr

nS)

Cis

tant

he sp

. ind

et. 1

1C

. sp_

00-2

132

CH

ILE.

IV: E

lqui

, rd

betw

. Mar

ques

a &

Con

doria

co k

m 1

2 N

of M

arqu

esa,

DQ

0902

12 /

Her

shko

vitz

00-

213

(CO

NC

). C

OM

MEN

T: R

esem

bles

gia

nt fo

rm o

fD

Q09

0147

Cis

tant

he a

rena

ria.

See

ds H

.

C. s

p_99

-910

2C

HIL

E. IV

: Cho

apa,

hill

side

S o

f Illa

pel,

Her

shko

vitz

99-

910

(CO

NC

).D

Q09

0201

/D

Q09

0136

CO

MM

ENT:

Res

embl

es g

iant

form

of C

ista

nthe

are

nari

a.

R

osul

atae

form

s (=

Cal

andr

inia

sect

.

R

osul

atae

Rei

che

sens

u R

eich

e 18

98)

Cis

tant

he c

epha

laph

ora

(I. M

. Joh

nst.)

Car

olin

ex H

ersh

k.ce

phal

apho

ra2

CH

ILE.

II: T

alta

l, km

114

0 of

Pan

amer

ican

Hw

y, H

ersh

kovi

tz 0

0-27

(CO

NC

).D

Q09

0230

/D

Q09

0165

Cist

anth

e coq

uim

bens

is (B

arné

oud)

coqu

imbe

nsis

1,2,

3C

HIL

E. IV

: Elq

ui, P

anam

eric

an H

wy

km 4

89, c

a. 1

5 km

N o

f mai

n jc

tD

Q09

0233

/C

arol

in ex

Her

shk.

in L

a Se

rena

, Her

shko

vitz

00-

155

(CO

NC

).D

Q09

0168

Cis

tant

he c

ymos

a (P

hil.)

Her

shk.

cym

osa_

00-1

092

CH

ILE.

III:

Hua

sco,

Pan

amer

ican

Hw

y be

tw. V

alle

nar &

Cop

iapó

ca.

DQ

0902

34 /

21 k

m N

of V

alle

nar,

Her

shko

vitz

00-

109

(CO

NC

). C

OM

MEN

T:D

Q09

0169

Peta

ls p

ink,

see

ds G

.

cym

osa_

00-1

382

CH

ILE.

III:

Hua

sco,

rd S

from

Fre

irina

ca.

20

km S

of F

reiri

na, H

ersh

kovi

tzD

Q09

0235

/00

-138

(CO

NC

). C

OM

MEN

T: P

etal

s pi

nk, s

eeds

G.

DQ

0901

70

cym

osa_

00-1

951,

2,3

CH

ILE.

III:

Hua

sco,

rd fr

om D

omey

ko to

Cal

eta

Sarc

o 2

km W

DQ

0902

37 /

of D

omey

ko, H

ersh

kovi

tz 0

0-19

5 (C

ON

C).

CO

MM

ENT:

Pet

als

DQ

0901

72pi

nk, s

eeds

G.

cym

osa_

00-3

91,2,

3C

HIL

E. II

: Tal

tal,

rd b

etw.

Tal

tal &

Pap

oso

ca. 2

0 km

N o

f Tal

tal,

DQ

0902

16 /

Her

shko

vitz

00-

39 (C

ON

C).

CO

MM

ENT:

Pet

als

yello

w, s

eeds

G.

DQ

0901

51

Gayana Bot. 63(1), 2006

60

Tabl

e I,

cont

inue

d

Gen

Ban

k se

quen

ceTa

xon

Acr

onym

Spec

imen

and

col

lect

ion

data

and

com

men

tary

acce

ssio

n(I

TS/y

cf3-

trnS

)

cym

osa_

02-1

381

CH

ILE.

III:

Hua

sco,

rd fr

om D

omey

ko to

Cal

eta

Sarc

o 5

km W

of

DQ

0902

36 /

Dom

eyko

, Her

shko

vitz

02-

138

(CO

NC

). C

OM

MEN

T: P

etal

s pi

nk,

DQ

0901

71se

eds

G.

cym

osa_

02-5

91,2,

3C

HIL

E. V

: Pet

orca

, Pan

amer

ican

Hw

y nr

Los

Mol

les,

40 k

m S

of L

os V

ilos,

DQ

0902

05 /

Her

shko

vitz

02-

59 (C

ON

C).

CO

MM

ENT:

Pet

als

whi

te, s

eeds

G.

DQ

0901

40

Cis

tant

he sp

. ind

et. 1

1, a

ff. C

ista

nthe

x cy

mos

a2C

HIL

E. II

I: H

uasc

o, rd

from

Dom

eyko

to C

alet

a Sa

rco

5 km

W o

f Dom

eyko

,D

Q09

0213

/cy

mos

aH

ersh

kovi

tz 0

0-20

7 (C

ON

C).

CO

MM

ENT:

The

pla

nts

cooc

cur

with

DQ

0901

48pi

nk-f

low

ered

C. c

ymos

a, b

ut th

e flo

wer

s are

whi

te, t

he st

ems a

nd le

aves

abou

t tw

ice

as la

rge,

15-

17 st

amen

s vs 5

, and

seed

s PT

vs G

.

Cista

nthe

long

iscap

a (B

arné

oud)

Car

olin

long

isca

pa_0

0-10

02 C

HIL

E. II

I: H

uasc

o, P

anam

eric

an H

wy

betw

. La S

eren

a & V

alle

nar c

a.D

Q09

0218

/ex

Her

shk.

69 k

m fr

om V

alle

nar,

Her

shko

vitz

00-

100

(CO

NC

). C

OM

MEN

T: S

eeds

G.

DQ

0901

53

long

isca

pa_0

0-10

62C

HIL

E. II

I: H

uasc

o, P

anam

eric

an H

wy

betw

. Val

lena

r & C

opia

pó c

a.D

Q09

0207

/21

km

N o

f Val

lena

r, H

ersh

kovi

tz 0

0-10

6 (C

ON

C).

CO

MM

ENT:

See

ds G

.D

Q09

0142

long

iscap

a_00

-236

CH

ILE.

IV: E

lqui

, Rut

a 41

ca.

36

km E

of V

icuñ

a, H

ersh

kovi

tz 0

0-23

6D

Q09

0221

/(C

ON

C).

DQ

0901

56

long

isca

pa_0

0-23

82C

HIL

E. IV

: Elq

ui, R

uta

41 c

a. 4

9 km

E o

f Vic

uña,

Her

shko

vitz

00-

238

DQ

0902

22 /

(CO

NC

). C

OM

MEN

T: S

eeds

PT.

DQ

0901

57

long

isca

pa_0

0-28

1,2,

3C

HIL

E. II

: Tal

tal,

km 1

140

of P

anam

eric

an H

wy,

bet

w. A

ntof

agas

taD

Q09

0217

/&

Tal

tal,

Her

shko

vitz

00-

28 (C

ON

C).

CO

MM

ENT:

See

ds G

.D

Q09

0152

long

isca

pa_0

0-71

1,2,

3C

HIL

E. II

: Tal

tal,

Pana

mer

ican

Hw

y ca

. 20

km S

E of

Tal

tal,

Her

shko

vitz

DQ

0902

09 /

00-7

1 (C

ON

C).

CO

MM

ENT:

See

ds G

.D

Q09

0144


61

Tabl

e I,

cont

inue

dG

enB

ank

sequ

ence

Taxo

nA

cron

ymSp

ecim

en a

nd c

olle

ctio

n da

ta a

nd c

omm

enta

ryac

cess

ion

(ITS

/ycf

3-tr

nS)

long

isca

pa_0

0-72

B2

CH

ILE.

III:

Cop

iapó

, Pan

amer

ican

Hw

y 10

km

N o

f Cal

dera

, Her

shko

vitz

DQ

0902

19 /

00-7

2B (C

ON

C).

CO

MM

ENT:

See

ds G

.D

Q09

0154

long

isca

pa_0

0-21

22C

HIL

E. IV

: Elq

ui, r

d be

tw. M

arqu

esa

& C

ondo

riaco

km

12

N o

f Mar

ques

a,D

Q09

0203

/H

ersh

kovi

tz 0

0-21

2 (C

ON

C).

CO

MM

ENT:

See

ds P

T.D

Q09

0138

long

isca

pa_0

1-04

2C

HIL

E. IV

: Elq

ui, C

ordi

llera

Doñ

a Ana

, Fun

do O

livie

res,

Her

shko

vitz

DQ

0902

06 /

01-0

4 (C

ON

C).

CO

MM

ENT:

See

ds P

T.D

Q09

0141

long

isca

pa_0

1-10

82C

HIL

E. IV

: Elq

ui, P

anam

eric

an H

wy

betw

. Los

Vilo

s & L

a Ser

ena c

a. 1

km

DQ

0902

10 /

N o

f Las

Tac

as jc

t, H

ersh

kovi

tz 0

1-10

8 (C

ON

C).

DQ

0901

45

long

isca

pa_0

2-78

2C

HIL

E. IV

: Elq

ui, P

anam

eric

an H

wy

betw

. La S

eren

a & V

alle

nar,

km 5

65, c

a.D

Q09

0225

/91

km

from

La

Sere

na, H

ersh

kovi

tz 0

2-78

(CO

NC

). C

OM

MEN

T: S

eeds

G.

DQ

0901

60

long

isca

pa_0

2-12

82C

HIL

E. II

I: C

opia

pó, r

d to

war

d B

arra

nqui

llas 1

km

S o

f Pue

rto V

iejo

,D

Q09

0223

/H

ersh

kovi

tz 0

2-12

8 (C

ON

C).

DQ

0901

58

Cis

tant

he m

ariti

ma

(Nut

t.)m

ariti

ma1

MEX

ICO

. Baj

a C

alifo

rnia

Nor

te: R

ebm

an 1

616

(SD

).D

Q09

0231

/C

arol

in ex

Her

shk.

DQ

0901

66

Cis

tant

he sp

. ind

et. 1

2, a

ff. C

alan

drin

iath

yrso

idea

_00-

157,2

,3C

HIL

E. IV

: Elq

ui, P

anam

eric

an H

wy

ca. 1

5 km

N o

f mai

n jc

t in

La S

eren

a,D

Q09

0228

/th

yrso

idea

Rei

che

Her

shko

vitz

00-

15 (C

ON

C).

7.D

Q09

0163

thyr

soid

ea_0

0-11

02C

HIL

E. II

I: H

uasc

o, P

anam

eric

an H

wy

betw

. Val

lena

r & C

opia

pó c

a.D

Q09

0226

/21

km

N o

f Val

lena

r, H

ersh

kovi

tz 0

0-11

0 (C

ON

C).

DQ

0901

61

thyr

soid

ea_0

0-17

22C

HIL

E. II

I: H

uasc

o, rd

from

Val

lena

r to

Junt

a Val

eria

na 4

0 km

from

ctr.

DQ

0902

27 /

of V

alle

nar,

Her

shko

vitz

00-

172

(CO

NC

).D

Q09

0162

Gayana Bot. 63(1), 2006

62

Tabl

e I,

cont

inue

d

Gen

Ban

k se

quen

ceTa

xon

Acr

onym

Spec

imen

and

col

lect

ion

data

and

com

men

tary

acce

ssio

n(I

TS/y

cf3-

trnS

)

thyr

soid

ea_0

0-21

62C

HIL

E. IV

: Elq

ui, r

d be

tw. M

arqu

esa

& C

ondo

riaco

km

12

N o

f Mar

ques

a,D

Q09

0229

/H

ersh

kovi

tz 0

0-21

6 (C

ON

C).

DQ

0901

64

Cis

tant

he sp

. ind

et. 1

3C

. sp_

00-1

392

CH

ILE.

III:

Hua

sco,

rd S

from

Fre

irina

ca.

20

km S

of F

reiri

na, H

ersh

kovi

tzD

Q09

0220

/00

-139

(CO

NC

). C

OM

MEN

T: T

he p

lant

has

the

erec

t hab

it an

d lin

ear l

vs.

DQ

0901

55of

C. t

hyrs

oide

a bu

t lar

ge fl

ower

s sim

ilar t

o C

. lon

gisc

apa.

See

ds P

T.

C. s

p_00

-160

2C

HIL

E. IV

: Elq

ui, P

anam

eric

an H

wy

ca. 1

5 km

N o

f mai

n jc

t in

La S

eren

a,D

Q09

0204

/H

ersh

kovi

tz 0

0-16

0 (C

ON

C).

CO

MM

ENT:

The

pla

nt h

as th

e er

ect h

abit

and

DQ

0901

39lin

ear l

vs. o

f C. t

hyrs

oide

a bu

t lar

ge fl

ower

s sim

ilar t

o C

. lon

gisc

apa.

See

ds G

.

C. s

p_00

_264

1,2,

3C

HIL

E. IV

: Elq

ui, r

d be

tw. T

ambi

llo &

Pan

amer

ican

Hw

y ca

. km

8 W

of

DQ

0902

00 /

jct w

/Rut

a 43

, Her

shko

vitz

00-

264

(CO

NC

). C

OM

MEN

T: T

he p

lant

has

the

DQ

0901

35er

ect h

abit

and

linea

r lvs

. of C

. thy

rsoi

dea

but l

arge

flow

ers s

imila

r to

C.

aren

aria

. See

ds P

T.

C. s

p_01

-109

1,2,

3C

HIL

E. IV

: Elq

ui, P

anam

eric

an H

wy

betw

. Los

Vilo

s & L

a Ser

ena c

a. 1

km

DQ

0902

08 /

N o

f Las

Tac

as jc

t, H

ersh

kovi

tz 0

1-10

9 (C

ON

C).

CO

MM

ENT:

Pla

nt s

tunt

ed,

DQ

0901

43po

ssib

ly a

stre

ssed

seed

ling.

C. s

p_02

-752

CH

ILE.

IV: E

lqui

, Pan

amer

ican

Hw

y be

tw. L

a Ser

ena &

Val

lena

r, km

565

, ca.

DQ

0902

24 /

91 k

m fr

om L

a Se

rena

, Her

shko

vitz

02-

75 (C

ON

C).

CO

MM

ENT:

The

pla

ntD

Q09

0159

has t

he e

rect

hab

it an

d lin

ear l

vs. o

f C. t

hyrs

oide

a bu

t lar

ge fl

ower

s sim

ilar t

o C

.lo

ngis

capa

. See

ds G

.

C. s

p_02

-802,

3C

HIL

E. IV

: Elq

ui, P

anam

eric

an H

wy

betw

. La S

eren

a & V

alle

nar,

ca. k

m 5

84,

DQ

0902

14 /

ca. 6

km

E o

f cre

st o

f Cue

sta

Pajo

nale

s, H

ersh

kovi

tz 0

2-80

(CO

NC

).D

Q09

0149

CO

MM

ENT:

Thi

s pl

ant r

esem

bles

a s

tunt

ed p

lant

of C

. sec

t. C

ista

nthe

, in

parti

cula

r in

its e

rect

, lea

fy h

abit

and

glau

cous

text

ure.

C. s

p_02

-145

2C

HIL

E. II

I: H

uasc

o, rd

from

Dom

eyko

to C

alet

a Sa

rco

ca. 4

km

W o

f jct

toD

Q09

0202

/


63

Tabl

e I,

cont

inue

d

Gen

Ban

k se

quen

ceTa

xon

Acr

onym

Spec

imen

and

col

lect

ion

data

and

com

men

tary

acce

ssio

n(I

TS/y

cf3-

trnS

)

Cal

eta

Cha

ñara

l, H

ersh

kovi

tz 0

2-14

5 (C

ON

C).

CO

MM

ENT:

The

pla

nt h

as th

eD

Q09

0137

erec

t hab

it an

d lin

ear l

vs. o

f C. t

hyrs

oide

a bu

t lar

ge fl

ower

s sim

ilar t

o C

.lo

ngis

capa

. See

ds P

T.

Cis

tant

he se

ct. P

hilip

piam

ra

(K

untz

e) H

ersh

k.C

ista

nthe

am

aran

toid

es (P

hil.)

Her

shk.

Am

aran

toid

es_0

0-46

2C

HIL

E. II

: Tal

tal,

alon

g rd

bet

w. T

alta

l & P

apos

o ca

. 20

km N

of T

alta

l,D

Q09

0379

/H

ersh

kovi

tz 0

0-46

(C

ON

C).

DQ

0903

45

Am

aran

toid

es_0

0-57

B2

CH

ILE.

II: T

alta

l, al

ong

rd b

etw.

Tal

tal &

Pap

oso

ca. 2

6 km

N o

f Tal

tal,

DQ

0903

80 /

Her

shko

vitz

00-

57b

(CO

NC

).D

Q09

0346

Cis

tant

he ce

losi

oide

s (Ph

il.) H

ersh

k.ce

losi

oide

s_00

-108

2C

HIL

E. II

I: H

uasc

o, P

anam

eric

an H

wy

betw

. Val

lena

r & C

opia

pó c

a.D

Q09

0377

/21

km

N o

f Val

lena

r, 17

Sep

200

0, H

ersh

kovi

tz 0

0-10

8 (C

ON

C).

DQ

0903

43

celo

sioi

des_

00-2

392

CH

ILE.

IV: E

lqui

, Rut

a 41

ca.

49

km E

of V

icuñ

a, H

ersh

kovi

tzD

Q09

0392

/00

-239

(CO

NC

).D

Q09

0352

celo

sioi

des_

00-8

32C

HIL

E. II

I: C

opia

pó, r

d fr

om P

aipo

te to

Die

go d

e Alm

agro

ca.

55

km N

DQ

0903

91 /

of P

aipo

te, H

ersh

kovi

tz 0

0-83

(CO

NC

).D

Q09

0351

celo

sioi

des_

02-1

472

CH

ILE.

III:

Hua

sco,

Pan

amer

ican

Hw

y be

tw. D

omey

ko &

La S

eren

a ca.

DQ

0903

88 /

3 km

S o

f Dom

eyko

, Her

shko

vitz

02-

147

(CO

NC

).D

Q09

0354

Cis

tant

he sp

. 14

aff.

Phili

ppia

mra

pach

yphy

lla_0

0-08

2C

HIL

E. II

: Tal

tal,

Pana

mer

ican

Hw

y km

110

3, b

etw.

Ant

ofag

asta

& T

alta

l,D

Q09

0393

/pa

chyp

hylla

(Phi

l.) K

untz

e.H

ersh

kovi

tz 0

0-08

(C

ON

C).

DQ

0903

59

pach

yphy

lla_0

0-86

2C

HIL

E. II

I: C

opia

pó, r

d fr

om P

aipo

te to

Die

go d

e Alm

agro

ca.

55

km N

of

DQ

0903

95 /

Paip

ote,

Her

shko

vitz

00-

86 (

CO

NC

).D

Q09

0361

Gayana Bot. 63(1), 2006

64

Tabl

e I,

cont

inue

dG

enB

ank

sequ

ence

Taxo

nA

cron

ymSp

ecim

en a

nd c

olle

ctio

n da

ta a

nd c

omm

enta

ryac

cess

ion

(ITS

/ycf

3-tr

nS)

C

ista

nthe

sect

ions

Am

aran

toid

es

or P

hilli

piam

ra in

cert

ae se

dis

Cis

tant

he sp

. ind

et. 1

5C

. sp_

00-1

02C

HIL

E. II

: Tal

tal,

Pana

mer

ican

Hw

y km

110

3, b

etw.

Ant

ofag

asta

& T

alta

l,D

Q09

0390

/H

ersh

kovi

tz 0

0-10

(CO

NC

). C

OM

MEN

T: P

lant

ste

rile,

ste

ms

spre

adin

g,D

Q09

0356

stiff

, yel

low

-gre

en, l

eave

s bro

adly

obl

ance

olat

e, su

rfac

e tex

ture

abra

sive

.

C. s

p_00

-112

CH

ILE.

II: T

alta

l, Pa

nam

eric

an H

wy

km 1

103,

bet

w. A

ntof

agas

ta &

Tal

tal,

DQ

0903

89 /

Her

shko

vitz

00-

11 (C

ON

C).

CO

MM

ENT:

Pla

nt s

teril

e, s

tem

s pr

ostra

te, l

ax,

DQ

0903

55m

agen

ta, l

eave

s spa

tula

te, n

otic

ibly

coo

l to

touc

h, su

rface

text

ure

smoo

th. T

hese

ster

ile p

lant

s coo

ccur

red

with

Her

shko

vitz

00-

10. S

uper

ficia

lly, t

hey

rese

mbl

edla

rge p

lant

s of P

ortu

laca

ole

race

a.Cl

ayto

nia

L.

C

layt

onia

sect

. Cau

dico

sae

A. G

ray

ex

P

oelln

itzC

layt

onia

meg

arhi

za (A

. Gra

y) S

. Wat

son

meg

arhi

za2,

3N

o vo

uche

r. C

ultiv

ated

pla

nt o

rigin

ally

colle

cted

in C

olor

ado,

USA

,D

Q09

0126

/de

term

ined

by

M. A

. Her

shko

vitz

.D

Q08

0655

Cla

yton

ia se

ct. C

layt

onia

Clay

toni

a vi

rigin

ica

virg

inic

a2,3

Vouc

her l

ost.

Orig

inal

dat

a: U

SA. M

aryl

and:

G. S

attle

r s.n

. 199

3, d

eter

min

edD

Q09

0125

/by

M. A

. Her

shko

vitz

DQ

0806

54

Lenz

ia P

hil.

L. ch

amae

pity

s Phi

l.ch

amae

pity

s2,3

CH

ILE.

III:

Hua

sco,

rd fr

om C

onay

to E

l Nev

ado

(Bar

rick

Min

e Pr

oyec

toD

Q09

0397

/Pa

scua

) at k

m 3

6, H

ersh

kovi

tz 0

1-40

(CO

NC

).D

Q09

0363

Lew

isia

Pur

shLe

wis

ia c

olum

bian

a (H

owel

l ex A

. Gra

y)ru

pico

la2

No

vouc

her.

Cul

tivat

ed, U

nive

rsity

of C

alifo

rnia

(Ber

kele

y) B

otan

ical

Gar

den

DQ

0901

23 /

B

. L. R

ob. v

ar. r

upic

ola

(Eng

lish)

acc.

89.

1410

, fro

m w

ild-c

olle

cted

pla

nt, S

. B. H

ogan

148

1, d

eter

min

ed b

y S.

DQ

0806

52

C. L

. Hitc

hc.

B. H

ogan

.


65

Tabl

e I,

cont

inue

dG

enB

ank

sequ

ence

Taxo

nA

cron

ymSp

ecim

en a

nd c

olle

ctio

n da

ta a

nd c

omm

enta

ryac

cess

ion

(ITS

/ycf

3-tr

nS)

Lew

isia

redi

viva

Pur

sh v

ar. m

inor

(Ryd

b.)

min

or2

USA

. Cal

iforn

ia: L

os A

ngel

es C

o., A

ngel

es N

atl.

Fore

st, H

ersh

kovi

tzD

Q09

0124

/M

unz

02-0

6 (C

ON

C).

DQ

0806

53Le

wis

iops

is R

. Gov

aerts

Lew

isio

psis

twee

dyi (

A. G

ray)

R. G

ovae

rtstw

eedy

i2,3

No

vouc

her.

Cul

tivat

ed, U

nive

rsity

of C

alifo

rnia

(Ber

kele

y) B

otan

ical

L780

89 /

Gar

den

acc.

89.

1401

, fro

m w

ild-c

olle

cted

pla

nt, S

. B. H

ogan

1145

,D

Q08

0651

dete

rmin

ed b

y S.

B. H

ogan

.

Mon

tia L

.M

ontia

cham

isso

i (Sp

reng

el) E

. Gre

ench

amis

soi2

USA

. Ore

gon:

Jack

son

Co.

, ca.

1 m

ile E

of L

inco

ln, H

ersh

kovi

tzD

Q09

0127

/02

-40

(CO

NC

).D

Q08

0656

Mon

tia h

owel

lii S

. Wat

son

how

ellii

2U

SA. C

alifo

rnia

: Trin

ity C

o., B

urnt

Ran

ch C

ampg

roun

d, H

ersh

kovi

tzD

Q09

0129

/02

-37

(CO

NC

).D

Q08

0658

Mon

tia li

near

is (H

ook.

) Gre

ene

linea

ris2

USA

. Mar

ylan

d: P

rince

Geo

rge’

s Co.

, Sm

ithso

nian

Inst

itutio

n M

useu

mD

Q09

0128

/Su

ppor

t Cen

ter,

Her

shko

vitz

s.n

. 10

May

198

6 (U

S)D

Q08

0657

Mon

tiops

is K

untz

e

Mon

tiops

is su

bg. D

iant

hoid

eae

(R

eich

e) D

. I. F

ord

Mon

tiops

is a

ndic

ola

(Gill

ies)

D. I

. For

dan

dico

la_2

4XI9

22,3

AR

GEN

TIN

A. M

endo

za: L

as H

eras

, wea

ther

sta

tion

at L

a C

umbr

e,D

Q09

0437

/Pe

ralta

& H

ersh

kovi

tz s.

n. 2

4 N

ov 1

992

(MER

L)D

Q09

0478

andi

cola

_98-

852

CH

ILE.

Met

ropo

litan

a: C

ordi

llera

, Val

le N

evad

o sk

i are

a, H

ersh

kovi

tzD

Q09

0438

/98

-85

(CO

NC

).D

Q09

0479

Mon

tiops

is ci

stiflo

ra (G

illie

s ex A

rn.)

cist

iflor

a2C

HIL

E. V

II: C

uric

ó, rd

from

Los

Que

ñes t

o Pa

so d

el P

lanc

hón

(Pas

o Ve

rger

a),

DQ

0904

39 /

D. I

. For

d1

rd k

m fr

om A

rgen

tina

bord

er, H

ersh

kovi

tz 9

9-54

6 (C

ON

C).

DQ

0904

80

Mon

tiops

is g

ayan

a (B

arné

oud)

D. I

. For

dga

yana

_99-

5172

CH

ILE.

VII

: Cur

icó,

rd fr

om L

os Q

ueñe

s to

Paso

del

Pla

nchó

n (P

aso

Verg

era)

,D

Q09

0434

/1

rd k

m fr

om A

rgen

tina

bord

er, H

ersh

kovi

tz 9

9-51

7 (C

ON

C).

DQ

0904

75

Gayana Bot. 63(1), 2006

66

Tabl

e I,

cont

inue

dG

enB

ank

sequ

ence

Taxo

nA

cron

ymSp

ecim

en a

nd c

olle

ctio

n da

ta a

nd c

omm

enta

ryac

cess

ion

(ITS

/ycf

3-tr

nS)

gaya

na_0

1-96

2C

HIL

E. V

III:

Ñub

le, T

erm

as d

e C

hillá

n, H

ersh

kovi

tz 0

1-96

.D

Q09

0435

/D

Q09

0476

gaya

na_4

242,

3C

ultiv

ated

, CH

ILE.

IX: e

x Fo

rd &

Ros

as 4

24b

(MO

).D

Q09

0436

/D

Q09

0477

Mon

tiops

is tr

icol

or (P

hil.)

D. I

. For

dtri

colo

r2C

HIL

E. M

etro

polit

ana:

Cor

dille

ra, V

alle

Nev

ado

hote

l par

king

area

,D

Q09

0440

/H

ersh

kovi

tz 9

8-58

(C

ON

C).

DQ

0904

81

M

ontio

psis

subg

. Mon

tiops

is

H

irsut

ae g

roup

(For

d 19

92)

Mon

tiops

is c

opia

pina

(Phi

l.) D

. I. F

ord

copi

apin

a_01

-352

CH

ILE.

III:

Hua

sco,

rd b

etw.

Con

ay &

El N

evad

o (B

arric

k M

ine,

Pro

yect

oD

Q09

0404

/Pa

scua

), ca

. km

35

S of

Con

ay, H

ersh

kovi

tz 0

1-35

(CO

NC

).D

Q09

0447

copi

apin

a_89

2871

CH

ILE.

IV: E

lqui

, Cor

dille

ra D

oña A

na, s

ki sl

ope

S of

Bañ

os d

el T

oro,

DQ

0904

11 /

Aran

cio

8928

7 (U

LS).

DQ

0904

53

copi

apin

a_92

1891

CH

ILE.

IV: E

lqui

, Cor

dille

ra D

oña A

na, V

ega P

iuqu

enes

, Squ

eo 9

2189

(ULS

)D

Q09

0412

/D

Q09

0454

Mon

tiops

is g

illie

sii (

Hoo

k. &

Arn

.) D

. I.

gilli

esii_

7381,

3A

RG

ENTI

NA

. Men

doza

: Las

Her

as, V

alle

Hor

cone

s, La

s Cue

vas-

Usp

alla

taD

Q09

0406

/Fo

rdrd

3 m

iles N

of R

uta

7, F

ord

& P

eral

ta 7

38 (M

O)

DQ

0904

49

gilli

esii_

99-5

421

CH

ILE.

VII

: Cur

icó,

rd fr

om L

os Q

ueñe

s to

Paso

del

Pla

nchó

n (P

aso

Verg

era)

,D

Q09

0413

/1

rd k

m fr

om A

rgen

tina

bord

er, H

ersh

kovi

tz 9

9-54

2 (C

ON

C).

DQ

0904

55

Mon

tiops

is po

tent

illio

ides

(Bar

néou

d)po

tent

illio

ides

2C

HIL

E. M

etro

polit

ana:

Cor

dille

ra, V

alle

Nev

ado

ski a

rea,

DQ

0904

07 /

D. I

. For

dH

ersh

kovi

tz 9

8-97

(C

ON

C).

DQ

0904

50

Mon

tiops

is se

rice

a (H

ook.

& A

rn.)

D. I

. For

dse

ricea

_12X

II03

1C

HIL

E. V

: Qui

llota

, Par

que

Nac

iona

l de

La C

ampa

na, s

ecto

r nea

r ent

ranc

e at

DQ

0904

08 /

Gra

nizo

, Rou

gier

s.n.

, 12

Dec

200

3 (n

o vo

uche

r).

DQ

0904

51


67

Tabl

e I,

cont

inue

dG

enB

ank

sequ

ence

Taxo

nA

cron

ymSp

ecim

en a

nd c

olle

ctio

n da

ta a

nd c

omm

enta

ryac

cess

ion

(ITS

/ycf

3-tr

nS)

Mon

tiops

is u

mbe

llata

(Rui

z &

Pav

.) D

. I. F

ord

umbe

llata

2C

ultiv

ated

, ex

Andr

ews &

Arc

hiba

ld 1

2535

(MO

).D

Q09

0405

/D

Q09

0448

Mon

tiops

is u

spal

late

nsis

(Phi

l.) D

. I. F

ord

uspa

llate

nsis

1C

HIL

E. IV

: Lim

ari,

Que

brad

a Lar

ga, S

side

of R

io L

os M

olle

s, Fo

rd &

DQ

0904

09,

Aran

cio

754

(MO

).D

Q09

0410

/D

Q09

0452

Con

dens

atae

gro

up (F

ord

1992

)M

ontio

psis

cap

itata

(Hoo

k. &

Arn

.) D

. I. F

ord

capi

tata

1,2,

3C

HIL

E. IV

: Elq

ui, r

d be

tw. H

urta

do &

Pab

elló

n 24

.4 k

m E

of H

urta

do,

DQ

0904

03 /

Her

shko

vitz

00-

257

(CO

NC

).D

Q09

0446

Mon

tiops

is g

lom

erat

a (P

hil.)

D. I

. For

dgl

omer

ata2,

3C

HIL

E. IV

: Elq

ui, s

lope

s nea

r Bañ

os d

el T

oro,

Her

nand

ez s.

n.D

Q09

0425

/Fe

b 20

03 (C

ON

C).

DQ

0904

67

Mon

tiops

is m

odes

ta (P

hil.)

D. I

. For

dm

odes

ta_0

2-94

1,2,

3C

HIL

E. II

I: H

uasc

o, rd

from

Val

lena

r to

Los M

orte

ros,

Her

shko

vitz

02-

94D

Q09

0420

/(C

ON

C).

CO

MM

ENT:

Thi

s spe

cies

has

nev

er b

een

colle

cted

bel

ow 3

000

mD

Q09

0462

nor b

efor

e la

te D

ecem

ber i

n R

egio

n II

I, no

r sym

patri

cally

with

M. p

arvi

flora

.

mod

esta

_II2

0032,

3C

HIL

E. IV

: Elq

ui, s

lope

s nea

r Bañ

os d

el T

oro,

Her

nand

ez s.

n. F

eb 2

003

DQ

0904

19 /

(CO

NC

).D

Q09

0461

Mon

tiops

is tr

ifida

(Hoo

k. &

Arn

.) D

. I. F

ord

trifid

a_00

-137

2C

HIL

E. II

I: H

uasc

o, c

a. 1

0.5

km S

from

Fre

irina

, Her

shko

vitz

00-

137

(CO

NC

). D

Q09

0426

/D

Q09

0468

trifid

a_00

-258

1,2,

3C

HIL

E. IV

: Elq

ui, r

d be

tw. V

icuñ

a & H

urta

do 2

2 km

S o

f Vic

uña,

Her

shko

vitz

DQ

0904

27 /

00-2

58 (C

ON

C).

DQ

0904

69

trifid

a_26

42,3

CH

ILE.

IV: E

lqui

, Min

a El R

omer

al, 3

km

from

Pan

amer

ican

Hw

y on

alte

rnat

eD

Q09

0428

/S

acce

ss rd

, For

d 26

4 (M

O).

DQ

0904

70

trifid

a_99

-898

1,2,

3C

HIL

E. IV

: Cho

apa,

rd b

etw.

Illa

pel &

Com

barb

alá,

Her

shko

vitz

99-

898

DQ

0904

29,

(CO

NC

).D

Q09

0430

/

Gayana Bot. 63(1), 2006

68

Tabl

e I,

cont

inue

d

Gen

Ban

k se

quen

ceTa

xon

Acr

onym

Spec

imen

and

col

lect

ion

data

and

com

men

tary

acce

ssio

n(I

TS/y

cf3-

trnS

)

DQ

0904

71,

DQ

0904

72

Mon

tiops

is tri

fida

x ca

pita

tax

trifid

a,2,3

CH

ILE.

IV: L

imar

í, R

ío C

ogat

í can

yon,

Her

shko

vitz

02-

152

(CO

NC

).D

Q09

0432

/C

OM

MEN

T: T

he m

orph

olog

y an

d el

evat

ion

corr

espo

nd w

ith th

ose

give

n by

DQ

0904

73Fo

rd (1

992)

for s

uch

inte

rspe

cific

hyb

rids.

Par

viflo

rae

grou

p (F

ord

1992

)M

ontio

psis

ber

tero

ana

(Phi

l.) D

. I. F

ord

berte

roan

a2,3

Vouc

her l

ost.

Orig

inal

dat

a: C

HIL

E. V

III:

Bío

-Bío

, Rd

betw

. Yum

bel &

Est

ació

nD

Q09

0398

/Yu

mbe

l at b

ridge

ove

r Río

Cla

ro, H

ersh

kovi

tz 9

2-11

, det

erm

ined

by

DQ

0904

41M

. A. H

ersh

kovi

tz.

Mon

tiops

is c

umin

gii (

Hoo

k. &

Arn

.) D

. I. F

ord

cum

ingi

i1C

HIL

E. I:

Par

inac

ota,

Veg

as d

e Caq

uena

, Ara

ncio

926

91 (U

LS).

DQ

0903

99 /

DQ

0904

42

Mon

tiops

is d

emis

sa (P

hil.)

D. I

. For

dde

mis

sa_0

0-21

02C

HIL

E. IV

: Elq

ui, P

anam

eric

an H

wy

betw

. La S

eren

a & V

alle

nar,

km 6

2 N

DQ

0904

00 /

of m

ain

jct i

n La

Ser

ena,

Her

shko

vitz

00-

210

(CO

NC

).D

Q09

0443

dem

issa

_02-

1431,

2,3

CH

ILE.

III:

Hua

sco,

rd fr

om D

omey

ko to

Cal

eta

Sarc

o ca

. 4 k

m W

of j

ct to

DQ

0904

01 /

Cal

eta

Cha

ñara

l, H

ersh

kovi

tz 0

2-14

3 (C

ON

C).

DQ

0904

44

Mon

tiops

is p

arvi

flora

(Phi

l.) D

. I. F

ord

parv

iflor

a_00

-184

1,2,

3C

HIL

E. II

I: H

uasc

o, rd

from

Val

lena

r to

Junt

a Val

eria

na 4

km

E o

f Con

ay,

DQ

0904

24 /

Her

shko

vitz

00-

184

(CO

NC

).D

Q09

0466

parv

iflor

a_00

-217

1,2,

3C

HIL

E. IV

: Elq

ui, r

d be

tw. M

arqu

esa

& C

ondo

riaco

,D

Q09

0433

/H

ersh

kovi

tz 0

0-21

7 (C

ON

C).

DQ

0904

74

parv

ifolia

_02-

921,

2,3

CH

ILE.

III:

Hua

sco,

rd fr

om V

alle

nar t

o Lo

s Mor

tero

s, H

ersh

kovi

tzD

Q09

0422

/02

-92

(CO

NC

).D

Q09

0464


69

Tabl

e I,

cont

inue

dG

enB

ank

sequ

ence

Taxo

nA

cron

ymSp

ecim

en a

nd c

olle

ctio

n da

ta a

nd c

omm

enta

ryac

cess

ion

(ITS

/ycf

3-tr

nS)

parv

ifolia

_02-

931,

2,3

CH

ILE.

III:

Hua

sco,

rd fr

om V

alle

nar t

o Lo

s Mor

tero

s, H

ersh

kovi

tzD

Q09

0423

/02

-93

(CO

NC

).D

Q09

0465

parv

ifolia

_02-

951,

2,3

CH

ILE.

III:

Hua

sco,

rd fr

om V

alle

nar t

o Lo

s Mor

tero

s, H

ersh

kovi

tzD

Q09

0421

/02

-95

(CO

NC

).D

Q09

0463

Mon

tiops

is ra

mos

issi

ma

(Hoo

k. &

Arn

.)ra

mos

issi

ma_

00-1

582

CH

ILE.

IV: E

lqui

, Pan

amer

ican

Hw

y be

tw. L

a Ser

ena &

Val

lena

r, km

62

ND

Q09

0416

/D

. I. F

ord

of m

ain

jct i

n La

Ser

ena,

Her

shko

vitz

00-

158

(CO

NC

).D

Q09

0458

ram

osis

sim

a_00

-211

2C

HIL

E. IV

: Elq

ui, P

anam

eric

an H

wy

betw

. La S

eren

a & V

alle

nar,

km 6

2 N

DQ

0904

15 /

of m

ain

jct i

n La

Ser

ena,

Her

shko

vitz

00-

211

(CO

NC

).D

Q09

0457

ram

osis

sim

a_C

HIL

E. IV

: Lim

arí,

rd b

etw

. Pun

itaqu

i & C

omba

rbal

á 36

km N

DQ

0904

14 /

02-1

48B

1,2,

3 o

f Com

barb

alá,

Her

shko

vitz

02-

148B

(CO

NC

).D

Q09

0456

ram

osis

sim

a_99

-900

2C

HIL

E. IV

: Cho

apa,

rd b

etw.

Illa

pel &

Com

barb

ala,

Her

shko

vitz

99-

900

DQ

0904

02 /

(CO

NC

).D

Q09

0445

ram

osiss

ima_

CH

ILE.

Met

ropo

litan

a: C

haca

buco

, rd

betw

. Pol

paic

o &

Tilt

il ca

. 9 k

m N

DQ

0904

18 /

99-9

651,

2,3

of P

olpa

ico,

Her

shko

vitz

99-

965

(CO

NC

).D

Q09

0460

ram

osiss

ima_

CH

ILE.

Met

ropo

litan

a: C

haca

buco

, Rd

betw

. Tilt

il &

Que

brad

a Alv

arad

oD

Q09

0417

/99

-973

1,2,

31

km W

of T

iltil,

Her

shko

vitz

99-

973

(CO

NC

).D

Q09

0459

Para

keel

ya H

ersh

k.Pa

rake

elya

lini

flora

(Fen

zl) H

ersh

k.lin

iflor

a1A

UST

RA

LIA

. Wes

tern

Aus

tral

ia: r

d fr

om M

andu

rah

to B

unbu

ry c

a. 1

4 m

iD

Q09

0130

/N

of j

ct w

/rd to

War

oona

, Deb

urh

3468

(RSA

)D

Q08

0659

Para

keel

ya p

tych

ospe

rma

(F. M

uell.

)pt

ycho

sper

ma2,

3C

ultiv

ated

, ex

J. G

. Wes

t 424

4 (C

AN

B)

L780

50 /

Her

shk.

DQ

0806

60

Gayana Bot. 63(1), 2006

70

Tabl

e I,

cont

inue

d

Gen

Ban

k se

quen

ceTa

xon

Acr

onym

Spec

imen

and

col

lect

ion

data

and

com

men

tary

acce

ssio

n(I

TS/y

cf3-

trnS

)

Phem

eran

thus

Raf

.Ph

emer

anth

us b

revi

foliu

s (To

rr.) H

ersh

k.br

evifo

lius2,

3N

o vo

uche

r. O

rigin

al d

ata:

Cul

tivat

ed, U

nive

rsity

of C

alifo

rnia

Bot

anic

al L

7803

8 /

Gar

den,

not

acce

ssio

ned,

orig

inal

colle

ctio

n, U

SA. A

rizo

na: C

ocon

ino

Co.

,D

Q08

0661

Mar

ble

Can

yon,

S. B

. Hog

an s.

n., d

eter

min

ed b

y S.

B. H

ogan

.

Talin

um A

dans

.Ta

linum

pan

icul

atum

(Jac

q.) G

aertn

.pa

nicu

latu

m2,

3N

o vo

uche

r. O

rigin

al d

ata

Cul

tivat

ed, U

SA. T

exas

: Bex

ar C

o., S

an A

nton

io,

L780

94 /

cour

tyar

d of

the

Men

ger H

otel

, Her

shko

vitz

s.n.

Aug

199

1, d

eter

min

ed b

yD

Q08

0662

M. A

. Her

shko

vitz

.


71

Prim

er n

ame

Sequ

ence

Com

men

t or

ref

eren

ce

ITS6

5’-T

TT

CT

TT

TC

CT

CC

GC

TTA

-3’

3’ (

reve

rse)

prim

er f

or d

oubl

e-st

rand

ed (

ds)

ampl

ifica

tion

of t

he I

TS r

egio

n; c

ompl

emen

tary

to

26S

rDN

A

ITS7

5’-G

AA

GG

AG

AA

GTC

GTA

AC

AA

G-3

’5’

(fo

rwar

d) p

rimer

for

ds

ampl

ifica

tion

of t

he I

TS r

egio

n; c

ompl

emen

tary

to

18S

rDN

A

ITS4

5’-T

CC

TC

CG

CT

TAT

TG

ATA

TG

C-3

’In

tern

al t

o IT

S6 p

rimer

and

use

d to

gen

erat

e IT

S re

gion

non

codi

ng s

trand

in

sing

le-s

trand

(ss

) PC

R a

nd

orse

quen

cing

of

codi

ng s

trand

ss

PCR

pro

duct

.

N18

L18

5’-A

AG

TCG

TAA

CA

AG

GTT

TC-3

’In

tern

al t

o IT

S7 p

rimer

and

use

d to

gen

erat

e IT

S re

gion

non

codi

ng s

trand

in

sing

le-s

trand

(ss

) PC

R a

ndfo

r se

quen

cing

of

codi

ng s

trand

ss

PCR

pro

duct

.

PORT

ITS4

R5’

-GG

TCG

CA

AC

GG

TTTG

TG-3

’Po

rtula

cace

ae-s

peci

fic p

rimer

use

d to

gen

erat

e IT

S re

gion

non

codi

ng s

trand

in

asym

met

ric P

CR

and

for

sequ

enci

ng o

f co

ding

stra

nd s

s PC

R p

rodu

ct.

This

prim

er w

as u

sefu

l fo

r D

NA

ext

ract

ions

fro

mhe

rbar

ium

spe

cim

ens,

whe

re p

ossi

ble

cont

amin

atio

n w

ith n

on-P

ortu

laca

ceae

DN

A w

as a

pro

blem

.

SP43

095F

5’-T

TT

CT

CC

TG

AA

GT

TG

TC

GG

AA

T-3’

5’ (

forw

ard)

prim

er u

sed

for

ds a

mpl

ifica

tion

of t

he c

pDN

A y

cf3-

trnS

int

erge

nic

spac

er.

SP43

122F

5’-A

TTG

GC

YAC

AAY

TGA

AA

AG

G-3

’Pr

imer

use

d fo

r ge

nera

tion

of t

he y

cf3-

trnS

non

codi

ng s

trand

in

ss P

CR

and

for

seq

uenc

ing

of c

odin

gst

rand

ss

PCR

pro

duct

.

SP44

097R

5’-A

TTC

GA

AC

CC

TCG

GTA

AA

CA

-3’

3’ (

reve

rse)

prim

er u

sed

for

ds P

CR

of

the

cpD

NA

ycf

3-tr

nS i

nter

geni

c sp

acer

.

TAB

LE II

. Am

plifi

cafio

n an

d se

quen

cing

prim

ers u

sed

in th

e pr

esen

t stu

dy.

TAB

LA II

. Par

tidor

es d

e am

plifi

caci

ón y

secu

enci

ació

n u

sada

s en

el p

rese

nte

estu

dio.

Gayana Bot. 63(1), 2006

72

TAB

LE II

I. Va

riatio

n in

alig

ned

wes

tern

Am

eric

an P

ortu

laca

ceae

ITS

& yc

f3-tr

nS se

quen

ces.

The t

able

lists

the n

umbe

r of v

aria

ble/

info

rmat

ive c

hara

cter

s and

info

rmat

ive i

ndel

char

acte

rs at

var

ious

phy

loge

netic

leve

ls. T

he n

umbe

r of i

nfor

mat

ive c

hara

cter

s is i

nclu

sive

of t

he n

umbe

r of i

nfor

mat

ive i

ndel

s. Th

e num

ber o

f inf

orm

ativ

e ind

els r

efer

s to

thos

e ind

els t

hat w

ere c

onsi

dere

d un

ambi

guou

sly

alig

nabl

e. A

dditi

on in

dels

occ

ur in

the d

ata.

The

bp

num

bers

incl

ude o

nly

varia

tion

at al

igne

d po

sitio

ns, e

.g.,

a reg

ion

of yc

f3-

trnS

was

una

ligna

ble

at th

e in

terg

ener

ic le

vel (

see

resu

lts),

henc

e va

riatio

n in

this

regi

on is

not

incl

uded

in th

e su

ms b

elow

. The

dat

a ar

e in

tend

ed p

rimar

ily fo

r com

paris

ons

betw

een

taxa

and

bet

wee

n da

ta se

ts.

TAB

LA II

I. Va

riaci

ón d

e las

secu

enci

as al

inea

das d

e ITS

& yc

f3-tr

nS d

e Por

tula

cace

ae d

e Am

éric

a occ

iden

tal.

Se in

dica

n el

núm

ero

de ca

ract

eres

var

iabl

es/in

form

ativ

os y

de

inse

rció

n/de

leci

ón in

form

ativ

os en

var

ios n

ivel

es fi

loge

nétic

os. E

l núm

ero

de ca

ract

eres

incl

uye e

l núm

ero

de in

serc

ione

s/de

leci

ones

info

rmat

ivas

. El n

úmer

o de

inse

rcio

nes/

dele

cion

es in

form

ativ

os se

refie

re a

aque

llos q

ue fu

eron

cons

ider

ados

no

ambi

guos

al al

inea

rse.

Inse

rcio

nes/d

elec

ione

s adi

cion

ales

apar

ecen

en lo

s dat

os. E

l núm

ero

pb in

cluy

esó

lo la

var

iaci

ón en

las p

osic

ione

s alin

eada

s, e.

g., u

na re

gión

de y

cf3-

trnS

no fu

e alin

eada

a ni

vel i

nter

gené

rico

(ver

resu

ltado

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73

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