Abstract Genus Paspalum L. comprises approx-

imately 400 species worldwide and about 220 in

Brazil. Paspalum is ecologically and economically

important, and has been very useful as pasture.

Traditionally, taxonomists use informal groups,

most of which described by Agnes Chase in 1929.

Some groups present a problematic circumscrip-

tion, this is the case of the Linearia and Notata

groups. This work uses a phylogenetic approach to

study these groups and related species. DNA

sequences from ITS of nuclear rRNA, from chlo-

roplast intergenic spacer psbA-trnH and chloro-

plast trnL intron were used to perform the

analyses. The informal groups studied were con-

sidered highly artificial, being the representatives

from several informal groups splitted throughout

the trees. Only a small core of species from Notata

group could be accepted as a good formal clade.

Keywords Notata Æ Linearia Æ Paspalum L. ÆPoaceae Æ phylogeny

Introduction

Genus Paspalum L. belongs to the grass family

(Poaceae) and presents annual and perennial

species, distributed around the tropics and sub-

tropics of both hemispheres. About 400 species

are described for this genus (Chase 1929; Oliveira

and Valls 2002). In Brazil, there are not studies

with detailed information about the total number

of species, nevertheless Valls and Pozzobon

(1987) estimated the occurrence of about 220

species.

Paspalum presents several species with poten-

tial forage value, species which are sometimes

known as weedy, other are used as ornamental

plants in green gardens, parks, soccer fields and one

species is used as cereal in India, P. scrobiculatum

L., called ‘‘Kodo’’ (Clayton and Renvoize 1986).

In Brazil, the genus is dominant in many native

herbaceous communities and constitutes the

major part of available forage. Paspalum notatum

Fluegge is the most common species of the genus in

Southern Brazil and is widely used as forage and as

an ornamental plant.

In 1929, Agnes Chase divided the species

occurring in North America into 25 groups,

without a formal taxonomic category (informal

L. Essi (&)Programa de Pos-graduacao em Botanica,Universidade Federal do Rio Grande do Sul, Av.Bento Goncalves, 9500, Campus do Vale da UFRGS,Predio 43433. Bairro Agronomia, Porto AlegreRS 91501-970, Brazile-mail: liliana_essi@yahoo.com.br

T. T. de Souza-ChiesDepartamento de Botanica, Programa dePos-graduacao em Botanica, Universidade Federal doRio Grande do Sul, Porto Alegre, RS 91501-970,Brazil

Genet Resour Crop Evol (2007) 54:779–791

DOI 10.1007/s10722-006-9148-7

123

ORIGINAL PAPER

Phylogeny of Linearia and Notata groups of Paspalum L.(Poaceae, Panicoideae, Paniceae) and related species

Liliana Essi Æ Tatiana Teixeira de Souza-Chies

Received: 5 December 2005 / Accepted: 27 March 2006 / Published online: 7 December 2006� Springer Science+Business Media, Inc. 2006

groups). These species were grouped according to

their habit, inflorescence type, spikelet form and

dimensions, pilosity and foliar width. These

informal groups proposed by Chase (1929)

became widely used by agrostologists, with the

addition of new groups proposed or adapted to

Chase’s groups, according to regional floras.

Actually, there is a lack of infrageneric divi-

sions to adequately describe the morphological

diversity in Paspalum, and the use of informal

groups attenuates this fact. However, the place-

ment of many taxa in these informal groups is still

controversial and some groups seem to be highly

artificial. A good example of this situation is seen

in the Linearia and Notata groups.

The Linearia group, as originally proposed

(Chase 1929), included only one species, P. lineare

Trin. Studies of other regional floras added to this

group species supposedly related to P. lineare

(Swallen 1967; Filgueiras 1982), or associated it to

species previously cited to other groups, especially

to Notata (Valls 1987; Oliveira and Valls 2002).

The Notata group (Chase 1929) initially in-

cluded five species. Barreto (1974) and Canto-

Dorow et al. (1996) expanded the original con-

cept, concerning the species from Rio Grande do

Sul state (Brazil), and accepting nine species.

Barreto (1974) also described a new group, El-

liptica, constituted only by P. ellipticum Doell, a

species included by Chase (unpubl. mss.) in

Lineares (=Linearia) and which, years later, Valls

(1987) pointed out as related to the Notata group.

Oliveira and Valls (2002) called attention to a

morphological continuity between Notata and

Linearia, emphasizing the condition of P. crom-

yorhizon Trin. ex Doell, P. ionanthum Chase and

P. dedeccae Quarın as species with intermediate

morphology between both groups. The same

authors also emphasized the morphological rela-

tionships among some Linearia species and spe-

cies belonging to other groups. For instance,

P. pallens Swallen was associated to P. vaginatum

Sw. (Disticha group), species from Recta group

were associated with Linearia, based on vegeta-

tive characters, and P. ovale Nees, included by

Chase in ‘‘Lineares’’ (unpubl. mss.), and its allies,

were suggested to be better placed in the Ovalia

group sensu Barreto (1974). Oliveira and Valls

(2002) emphasized the heterogeneity of the

Linearia group and the possible obtention of a

single more homogeneous group, if the discrepant

species, such as P. pallens, and species which

could be included in other groups, like P. ovale,

were excluded. The concept of a single group

including Linearia and Notata was adopted by

Zuloaga et al. (2004) in their recent monograph

on the Notata group. In this work, the authors

accept a group called Notata including major

species of Linearia according to the concept of

Oliveira and Valls (2002), but maintaining

P. pallens in this species complex.

The situations mentioned above illustrate

several controversies concerning the circum-

scription of Linearia and Notata groups, and their

monophyly is questioned. Table 1 shows a syn-

thesis of the placement of the species analyzed in

the present work, in informal groups, as proposed

by different authors.

Aliscioni (2002) evaluated the monophyly of

many informal groups of Paspalum, including a

small number of species belonging to Notata and

Linearia. This study was based on the anatomy of

leaves and exomorphological characters, and the

author concluded that major informal groups are

artificial, including Notata and Linearia. Consid-

ering the controversies around morphological

characters, many of them highly affected by the

environment or differentially classified by every

author, a contribution with characters of a dif-

ferent nature is needed.

Different DNA fragments have been used as

source of characters for phylogenetic analysis.

Outstanding among them are the Internal Tran-

scribed Spacers (ITS) located between the nuclear

rRNA genes, the chloroplast spacer psbA-trnH

and the chloroplast intron from trnL gene. The

first one has been widely used in plant phyloge-

nies, due to its high nucleotide substitution rates

(Baldwin 1992; Baldwin et al. 1995) and to its

power to elucidate relationships at the infragen-

eric level in plants of several families, including

Poaceae (Grebenstein et al. 1998; Catalan and

Olmstead 2000; Guo et al. 2002; Baumel et al.

2002). The second fragment was pointed out by

Sang et al. (1997) as phylogenetically informative,

and in Paspalum presented an adequate length for

direct sequencing (about 600 base pairs). The last

fragment has also been widely used to elucidate

780 Genet Resour Crop Evol (2007) 54:779–791

123

phylogenetic relationships among taxa from sev-

eral plant families (Stedje 1998; Doust and Kellog

2002; Goldblatt et al. 2002). The chloroplast

fragments are generally more conserved than ITS

and, for this reason, they have usually been com-

bined with other DNA fragments and/or with

characters of another nature.

In this work, a molecular approach was used in

order to evaluate the monophyly of Linearia,

Notata and related groups such as Disticha, Recta

and Ovalia, and to establish the phylogenetic

relationships among Paspalum species that be-

long to these groups, looking for clades in this

complex that could be assigned a formal category.

The data were evaluated using three methods of

phylogenetic inference: unweighted parsimony

(maximum parsimony or MP), maximum likeli-

hood (ML) and Bayesian estimation (B), as de-

scribed bellow.

Materials and methods

Plant material and laboratory methods

The 31 species included in this study are listed in

Table 2. Five Axonopus species constituted the

outgroup, and four Paspalum species belonging to

Table 1 Placement of the Paspalum species included in this work in the informal groups, according to different authors.

Species Informal groups according to the literature

P. almum Chase Notata (Chase 1933 ; Barreto 1974; Canto-Dorow et al. 1996), Alma(Quarın 1974; Valls 1987; Zuloaga et al. 2004)

P. barretoi Canto-Dorow, Valls etLonghi-Wagner

Notata (Canto-Dorow et al. 1995)

P. conduplicatum Canto-Dorow, Vallset Longhi-Wagner

Notata (Canto-Dorow et al. 1995; Zuloaga et al. 2004)

P. cromyorhizon Trin. ex Doell Notata (Barreto 1974; Canto-Dorow et al. 1996; Zuloaga et al. 2004)P. dedeccae Quarın Linearia (Oliveira and Valls 2002), Notata (Zuloaga et al. 2004)P. ellipticum Doell Linearia (as Lineares in Chase, unpublished; Oliveira and Valls 2002),

Elliptica (Barreto 1974), Notata (Zuloaga et al. 2004)P. equitans Mez Fasciculata (Barreto 1974)P. filifolium Nees ex Steud. Linearia (Oliveira and Valls 2002), Notata (Zuloaga et al. 2004)P. flaccidum Nees Recta (Chase, unpubl. mss.; Zuloaga et al. 2004)P. intermedium Munro ex Morong et

BrittonQuadrifaria (Barreto 1974)

P. ionanthum Chase Notata (Canto-Dorow et al. 1996; Zuloaga et al. 2004)P. juergensii Hack. Paniculata (Barreto 1974)P. lineare Trin. Linearia (Chase 1929; Oliveira and Valls 2002), Notata (Zuloaga et al.

2004)P. maculosum Trin. Notata (Barreto 1974; Canto-Dorow et al. 1996; Zuloaga et al. 2004),

Maculosa (Chase, unpubl. mss.)P. minus E. Fourn. Notata (Chase 1929; Barreto 1974; Canto-Dorow et al. 1996; Zuloaga et al.

2004)P. notatum Fluegge Notata (Chase 1929; Barreto 1974; Canto-Dorow et al. 1996; Zuloaga et al.

2004)P. nummularium Chase ex Sendulski et

A. G. BurmanNotata (Zuloaga et al. 2004)

P. ovale Nees ex Steud. Linearia (as Lineares in Chase, unpubl. mss.), Ovalia (Barreto 1974;Zuloaga et al. 2004)

P. pallens Swallen Linearia (Swallen 1967; Oliveira and Valls 2002), Notata (Zuloaga et al.2004)

P. pauciciliatum (Parodi) Herter Dilatata (Valls and Pozzobon 1987) Livida (Barreto 1974)P. pumilum Nees Notata (Canto-Dorow et al. 1996; Zuloaga et al. 2004)P. ramboi Barreto Notata (Barreto 1974; Canto-Dorow et al. 1996; Zuloaga et al. 2004)P. subciliatum Chase Notata (Chase 1929; Zuloaga et al. 2004)P. urvillei Steud. Dilatata (Barreto 1974)P. vaginatum Sw. Disticha (Chase 1929; Barreto 1974)

Genet Resour Crop Evol (2007) 54:779–791 781

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Genet Resour Crop Evol (2007) 54:779–791 783

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other informal groups not related to the complex

were arbitrarily chosen to test the Linearia/Notata

monophyly hypothesis. These species are consid-

ered well defined in Chase’s groups until the

present, clearly distinguished from the rest of

the ingroup, but not far enough to constitute

the outgroup: P. urvillei Steud., P. pauciciliatum

(Parodi) Herter, P. intermedium Munro ex

Morong & Britton and P. juergensii Hack. Genus

Axonopus was chosen to form the outgroup

because of its close morphological relationship

with Paspalum and, at the same time, because

of its clear and well-defined circumscription,

compared with the ingroup species. From the 21

species cited by Zuloaga et al. (2004) to Notata

group, 16 are included in this study, and other six

species belonging to closely related groups, or al-

ready cited by other authors to Notata or Linearia,

are also included. The sampling covered the main

controversies concerning the circumscription of

Linearia and Notata cited by Oliveira and Valls

(2002). All species included occur in Brazil.

The total DNA was isolated following a modi-

fied CTAB method (Doyle and Doyle 1987), from

dried or herbarium material. Voucher information

are listed in Table 2. The fragments were ampli-

fied using the primers described by Desfeux and

Lejeune (1996) to ITS, primers psbAF and trnHR

described by Sang et al. (1997) to psbA-trnH

spacer, and the primers C and D described by

Taberlet et al. (1991) to chloroplast intron from

trnL gene. The Polymerase Chain Reaction

(PCR) was performed following the recommen-

dations of the authors, with minor modifications.

The amplified material was purified with enzy-

matic pre-treatment (Shrimp alkaline phospha-

tase and exonuclease I, Amersham Biosciences),

and directly sequenced using the same primers

from PCR to chloroplast fragments and internal

primers to the ITS, also described by Desfeux and

Lejeune (1996). In the case of ITS, some of the

species were manually sequenced with chain ter-

minators labeled with 33P and Kit ‘‘Thermo

Sequenase’’. The other fragments and the ITS

region from the rest of the sampling were

sequenced using the automatic sequencers ‘‘ABI

Prism 310’’ (Perkin Elmer), ‘‘Mega Bace 1000’’

(Amersham-Pharmacia-Biotech) or ‘‘ABI Prism

3100’’ (Applied Biosystems). The ITS fragments

were sequenced at least twice, and the chloroplast

fragments were sequenced with forward and

backward primers. Ambiguous bases in the eletr-

opherograms were coded as ‘‘N’’. The sequences

obtained were stored in the GENBANK (http://

www.ncbi.nlm.nih.gov). All fragments were

sequenced to all species included in the study,

except the psbA-trnH spacer, which is not avail-

able for P. barretoi Canto-Dorow, Valls and

Longhi-Wagner and P. ovale.

Data analysis

The sequence alignments were made using the

software Clustal·1.81 (Thompson et al. 1997) and

manually refined with the help of the software

GeneDoc (Nicholas et al. 1997). The phyloge-

netic analyses were performed with the softwares

PAUP*, version 4.0b10 (Swofford 2002) and

Mr. Bayes, version 3 (Huesenbeck and Ronquist

2001). The analyses were made considering a

nuclear and a chloroplast data matrix (ITS or

psbA-trnH and trnL). ITS1 and ITS2 were always

analyzed together, as a single data collection. The

phylogenetic trees were calculated using three

methods: Maximum Parsimony (MP), Bayesian

(B) and Maximum Likelihood (ML). MP and ML

were performed with the PAUP* software, and B

was performed with Mr. Bayes software.

For MP analysis, heuristic searches with TBR

(tree bisection-reconnection), MULPARS option

and 1000 random-addition replications of heuristic

search and simple taxon addition, were used to all

trees saved option. The MAXTREES option was

selected, with auto-increase of 100. In the case of

the chloroplast matrix, the MAXTREES option

was set as 5000 trees due to the high number of

equally parsimonious trees. The relative propor-

tion of homoplasious data in the MP analysis was

estimated through Consistency and Retention

Indexes (CI and RI) available at PAUP* tools.

The heterogeneity among markers was assessed

using the partition homogeneity test (Farris et al.

1995), with 100 interactions, as implemented in

PAUP*. The G1 statistics (Hillis 1991) were

calculated through the generation of 10,000 ran-

dom trees by PAUP*. The tree-score frequency

distributions were ordenized by PAUP* as histo-

grams with 20 classes, showing the values to G1.

784 Genet Resour Crop Evol (2007) 54:779–791

123

Asymmetric histograms left inclinated and nega-

tive G1 values indicate that the trees generated

are unlikely to be produced by random, what

means that the fragments evaluated are phyloge-

netically informative. The Bootstrap test (Fel-

senstein 1985) was also performed with 1000

replications, heuristic search and simple taxon

addition, saving only the best trees, and the Bre-

mer support (Bremer 1988) was performed with

the Autodecay software, version 4.0.2’68k (Eri-

ksson 1999).

Nucleotide substitution models for ML and B

were chosen according to the best models selected

using the LTR criterion by software MODEL-

TEST 3.06 (Posada and Crandall 1998)-model

K80 + G to ITS matrix, model HKY85 + G to

chloroplast matrix and model TrN + I + G to the

combined matrix. ML analysis was performed with

PAUP* through heuristic search, as-is option,

TBR branch swapping and MULPARS option. B

analysis was performed with Mr. Bayes, through

1,000,000 generations.

Results and discussion

The ITS1 sequences presented lengths from

208 bp for Paspalum pallens and Axonopus

compressus (Sw.) P. Beauv., to 220 bp for P.

ionanthum and P. dedeccae, and the ITS2 length

varied between 207 bp for P. barretoi, and 220 bp

for P. cromyorhizon and A. brasiliensis (Spreng.)

Kuhlm. The total matrix length was 502 bp. When

single gaps were treated as missing data, the

matrix presented 269 conserved characters, 106

variable parsimony non-informative (autapomor-

phies) and 127 characters that were variable and

parsimony-informative. Considering gaps as fifth

base, the number of parsimony-informative

characters grew to 185. The average CG content

was 0.58.

The psbA-trnH spacer presented lengths vary-

ing from 534 bp for A. compressus, to 547 bp for

P. urvillei, and the CG content was 0.37 on the

average, similar to the CG content of the trnL

intron, which presented an average content of 0.34.

This intron presented lengths from 565 bp for

P. filifolium Nees ex Steud., to 590 bp for A. siccus

(Nees) Kuhlm. The whole chloroplast dataset was

1176 bp long, but with a low number of parsimony-

informative characters, 57 characters treating gaps

as fifth state character, and only 36 considering

gaps as missing data. The number of variable non-

informative characters was 142, treating gaps as

fifth base, and 64 treating gaps as missing data,

showing a high percentage of autapomorphic data.

The G1 statistic (Hillis 1991) for ITS matrix

was )1.635807, and for chloroplast matrix it was

)1.821921, presenting a phylogenetic signal good

enough for both datasets, although other statisti-

cal tests presented above pointed to a high degree

of conservation for the chloroplast matrix.

The measure of incongruence between the two

matrices, obtained through the Partition Homo-

geneity Test, rejected the null hypothesis, with a

strong significance level (P value = 0.01). This

clear incongruence between the chloroplast and

the nuclear dataset supported the decision to not

proceed with a combined analysis.

Statistics and phylogenetic trees

The phylogenetic trees inferred through ITS

(Fig. 1) and chloroplast (Fig. 2) showed quite

different groups and statistical values. The chlo-

roplast matrix provided a great number of possible

trees, and it was necessary to set a maximum

number of trees to be saved in MP analysis (5000).

The number of equally parsimonious trees was

always high for the chloroplast dataset, resulting

in highly polytomized consensus trees. It is also

interesting to take a look at the index used to

estimate homoplasy level. The chloroplast matrix

showed the lowest level of homoplasious data.

When treating single gaps as missing data,

CI and RI were the best (CI = 0.91, CI excluding

non-informative characters = 0.81, RI = 0.93),

but when gaps were included as fifth base,

the number presented a significant decrease

(CI = 0.85, CI excluding non-informative charac-

ters = 0.65, RI = 0.80). The index for ITS also

presented a significant decrease when including

gaps (from an RI = 0.71 to an RI = 0.60, for

instance), denoting that the MP analysis including

gaps could be not reliable enough. For this reason,

only trees produced with single gaps treated as

missing data are shown, although some results

with gaps as fifth base are commented.

Genet Resour Crop Evol (2007) 54:779–791 785

123

The trees inferred though ML presented

topologies similar to the B analyses, with minor

differences. In general, the groups supported by

bootstrap in MP analysis appeared in all other

phylogenetic analyses of the same dataset. Groups

not supported appeared in different positions

according to the phylogenetic method performed.

Groups that are not supported by any statistical

test and with an unstable position are not going to

be emphasized during the discussion. For instance,

it is possible to take a look at the behavior of

P. pallens in the trees. Considering the ITS matrix

and MP analyses, this species is placed within a

Notata clade, if treating gaps as fifth base. Treat-

ing gaps as missing data, its position changed, and

it was placed with P. almum Chase, as it can be

seen in Fig. 1. But considering a chloroplast ma-

trix and ML analysis, it was placed at the base of

the clade constituted by P. dedeccae, P. lineare, P.

notatum and P. vaginatum. Considering the

instability and lack of support of these relation-

ships, there is no reliable conclusion about its

phylogenetic relationships.

The position of P. chromyorhizon is always far

enough from P. notatum, against the possibility

of being considered a variety of P. notatum, as

Fig. 1 Consensus tree (majority rule values above thebranches) from 917 most parsimonious trees, with 442steps, inferred through ITS matrix, gaps treated as missingdata. Matrix with 127 parsimony-informative characters.

Support values included under the branches. D = decayvalues. B = bootstrap values. CI = 0.71, CI excluding non-informative characters = 0.60, IR = 0.70

786 Genet Resour Crop Evol (2007) 54:779–791

123

accepted by Smith and Wasshausen (1982). Also,

P. dedeccae, P. filifolium and P. ionanthum,

indicated by Oliveira and Valls (2002) as mor-

phological links between Notata and Linearia

groups, due to the continuity of their leaf shapes,

appeared always in distinct clades, showing that

these morphological affinities do not have the

same pattern at the molecular level.

From the ITS clades (Fig. 1) as a whole, it is

possible to highlight three structured clades. Clade

1 includes only species from the Notata group,

plus P. flaccidum Nees. Clade 2 includes species

from many informal groups, especially Linearia.

The clade 3 groups species representing the Con-

trol Group plus P. ionanthum and P. ovale.

Observing the distribution of the informal groups

in the tree, it can be seen that there is no mono-

phyletic group corresponding to Notata, Linearia

or both together, although a small part of Notata

could be considered monophyletic, when it in-

cludes P. flaccidum. This behavior agrees with the

work of Ferreira and Souza-Chies (2005), based

on RFLP data, but not with the total evidence

analysis including morphological data, which

Fig. 2 Consensus tree (majority rule values above thebranches) from 5000 most parsimonious trees, with 117steps, inferred through chloroplast matrix, gaps treated asmissing data. Matrix with 36 parsimony-informative

characters. Support values included under the branches.D = decay values. B = bootstrap values. CI = 0.91, CIexcluding non-informative characters = 0.81, IR = 0.93

Genet Resour Crop Evol (2007) 54:779–791 787

123

support the hypothesis of a single group formed by

Linearia plus Notata together (Zuloaga et al.

2004; Souza-Chies et al. 2006). Excluding P. flac-

cidum, the seven other species are in fact very

similar morphologically, so the clade maybe de-

serves some formal treatment.

Clade 2 is also very frequent in the analyses,

especially the subclade including P. equitans Mez,

P. ramboi Barreto and P. filifolium. The two

samples of P. ellipticum represent two distinct

biotypes, previously considered as different spe-

cies, but recently made synonymous (Oliveira and

Valls 2002). They are closely related, but curi-

ously do not present bootstrap support.

Clade 3, formed mostly by the species of other

informal groups morphogically well resolved, also

includes one species from the Ovale group and

another from the Notata group, showing that

neither species of those groups are far enough

from the complex, from the molecular standpoint.

In the chloroplast analysis (Fig. 2), other dis-

tinct clades appear. Clade B links P. barretoi and

P. nummularium Chase ex Sendulski et A.G.

Burman, both belonging to clade 1 from ITS

analysis, but not linked together in this clade.

Although their ITS sequences are very distinct,

their trnL intron sequences are almost identical,

differing in only one base pair. Clade D links

species not linked by ITS: P. dedeccae, P. lineare,

P. vaginatum and P. notatum. They belong to

different groups proposed by Chase and are

morphologically very distinct. Clade F links P.

subciliatum Chase and P. maculosum Trin. Both

belong to Notata, although P. maculosum pre-

sented an uncertain position in the ITS analysis

and Chase (unpublish. mss.) preferred to transfer

this species to the new Maculosa group.

The species P. almum appeared always far

from other Notata species. This species differs

from almost all species of Paspalum concerning

its chromosome number, which has a basic num-

ber of six instead of 10. On the other hand, Souza-

Chies et al. (2006) presented this species included

in the Linearia + Notata clade, a relationship

probably due mostly to the exomorphological

data.

Many authors have published chromosome

counts for species from several groups of Pas-

palum (e.g. Quarın and Burson 1983, 1991;

Quarın et al. 1984; Quarın and Norrmann 1987;

Norrmann et al. 1989; Pozzobon and Valls 1997;

Takayama et al. 1998; Pozzobon et al. 2000; and

Pagliarini et al. 2001), pointing out the occur-

rence of diploid, triploid, tetraploid, pentaploid,

hexaploid and also octoploid species. Many of

these species are possibly of hybrid origin, such as

the case of P. dilatatum Poir., which presents

three genomes, II, JJ and XX, that could be

donated by plants belonging to different groups,

like Quadrifaria and Paniculata (Casa et al. 2002,

Miz and Souza-Chies 2006). The occurrence of

hybridization in the genus can be an element that

makes it difficult to establish phylogenetic rela-

tionships. Natural hybrids can bring together

species that actually have a very different phy-

logeny. When a chloroplast fragment has unipa-

rental origin (as in the case of major

angiosperms), and it presents enough informative

sites, it is possible to recognize hybridization

cases, by comparisons between phylogenies pro-

duced by another molecular marker which is

inherited only from the opposite parent or has

biparental origin.

To avoid the distortions that could be pro-

duced by possible hybrid species, diploid speci-

mens were preferred for this work, whenever

possible (for chromosome counts, see Table 1).

Any further doubts could be checked contrasting

the phylogenies from chloroplast and nuclear

sources.

Differences between the ITS and chloroplast

trees topologies could be attributed to the effect

of some species of hybrid origin, but the poor

resolution of the chloroplast trees does not allow

for a definitive conclusion about the possible hy-

brids and parents, although some comments could

be made concerning chromosome numbers.

Paspalum ramboi, placed at clade 1 in the ITS

tree, which groups major Notata specimens,

appears linked to P. conduplicatum (clade C) in

the chloroplast tree. Both are the only hexaploid

Paspalum species included in the analyses. Clade

E in the chloroplast tree links plants from dif-

ferent clades of the ITS analysis, which, however,

share the same chromosome number: 2n = 20.

The clades A, F and G link species with different

chromosome numbers. Further details on the

relationships between major species included in

788 Genet Resour Crop Evol (2007) 54:779–791

123

clades A and G are being studied in our labora-

tory.

Conclusion

The three phylogenetic methods produced similar

trees, if only branches with good support or high

frequency in majority rule consensus are consid-

ered. Polytomized regions in strict consensus

presented different topologies when shown in

majority consensus from MP, B and ML. Since

MP trees were submitted to more statistical tests

than ML and B, they were the first choice to

synthesize the results obtained.

From the clades with high frequency, the most

similar to a previously circumscribed informal

group was clade 1 from ITS analyses, which also

occurred in combined analyses (not shown) but

excluding P. pallens. Another clade whose species

were already pointed out as related, if excluding P.

ovale, is clade 3. In general, representatives from

several informal groups are spread throughout the

trees, and only clade 1 (Notata group sensu stricto)

could be accepted as a good formal clade, due to its

high frequency in different analyses, to the good

support of some pairs, and to the morphological

similarities already observed by other authors.

The genus as a whole has complex reproduc-

tion and hybridization mechanisms, with sexual

and or apomitic plants, and this intricate web is

certainly being reflected in these molecular

markers. The unresolved groups can be better

studied in future analyses, considering the genus

as a whole, which is pointed out as paraphyletic

(Duvall et al. 2001; Giussani et al. 2001; Aliscioni

2002). The addition of species from other mor-

phological groups and the increase of informative

characters (molecular, morphological and cyto-

genetic) should improve the resolution and sup-

port of such groups, and bring new insights on the

relationships among species of this rich and

complex genus.

Acknowledgements We thank Drs. Jose F. M. Valls andRegina C. de Oliveira for the plant accessions and infor-mation supplied, Drs. Sandro Bonatto, Maurıcio R. Bogoand Giancarlo Pasquali for the use of sequencing facilities,and anonymous reviewers for suggestions. We also thankthe Genetics Department of Universidade Federal do Rio

Grande do Sul as a whole, for the welcome and theexchange of experiences, with a special thank to Dr.Alessandra Selbach. FAPERGS and CNPq, Brazil, sup-ported this work.

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