Internal transcribed spacer sequence phylogeny of Crambe L. (Brassicaceae): molecular data reveal two Old World disjunctions

Crambe L. (Brassicaceae) is an Old World genus with a disjunct distribution among four major centers of species diversity. A phylogenetic analysis of nucleotide sequences of the internal transcribed spacers (ITS) of the nuclear ribosomal repeat was conducted with 27 species of Crambe and 18 related genera. Cladistic analyses using weighted and unweighted parsimony support Crambe as a monophyletic genus with three major lineages. The first comprises those taxa endemic to the Macaronesian archipelagos. Taxa with a predominant Mediterranean distribution form the second assemblage, and a disjunction between east Africa (C. abyssinica) and the Mediterranean (C. hispanica) occurs in this clade. The third lineage includes all Eurosiberian-Asian taxa and C. kilimandscharica, a species from the highlands of east Africa. A basal biogeographic split between east Africa and Eurasia is present in the third clade. The patterns of relationships in the ITS tree are concordant with known climatic events in northern Africa and southwestern Asia since the middle Miocene. The ITS trees are congruent with the current sectional classification except for a few members of sections Crambe, Leptocrambe, and Orientecrambe (C. cordifolia, C. endentula, C. kilimandscharica, and C. kotschyana). Low levels of support in the basal branches do not allow resolution of which genera of the subtribes Raphaniae or Brassicinae are sister to Crambe. Both subtribes appear to be highly polyphyletic in the ITS trees.     

Phylogenetic relationships of Combretoideae (Combretaceae) inferred from plastid,nuclear gene and spacer sequences

Phylogenetic relationships of the subfamily Combretoideae (Combretaceae) were studied based on DNA sequences of nuclear ribosomal internal transcribed spacer (ITS) regions, the plastid rbcL gene and the intergenic spacer between the psaA and ycf3 genes (PY-IGS), including 16 species of eight genera within two traditional tribes of Combretoideae, and two species of the subfamily Strephonematoideae of Combretaceae as outgroups. Phylogenetic trees based on the three data sets (ITS, rbcL, and PY-IGS) were generated by using maximum parsimony (MP) and maximum likelihood (ML) analyses. Partition-homogeneity tests indicated that the three data sets and the combined data set are homogeneous. In the combined phylogenetic trees, all ingroup taxa are divided into two main clades, which correspond to the two tribes Laguncularieae and Combreteae. In the Laguncularieae clade, two mangrove genera, Lumnitzera and Laguncularia, are shown to be sister taxa. In the tribe Combreteae, two major clades can be classified: one includes three genera Quisqualis, Combretum and Calycopteris, within which the monophyly of the tribe Combreteae sensu Engler and Diels including Quisqualis and Combretum is strongly supported, and this monophyly is then sister to the monotypic genus Calycopteris; another major clade includes three genera Anogeissus, Terminalia and Conocarpus. There is no support for the monophyly of Terminalia as it forms a polytomy with Anogeissus. This clade is sister to Conocarpus. Electronic Publication     

<Emphasis Type="Italic">Welwitschiella</Emphasis> is a member of the African subtribe Grangeinae (Asteraceae Astereae): a new phylogenetic position based on <Emphasis Type="Italic">ndh</Emphasis>F and ITS sequence data

The African genus Welwitschiella has traditionally been placed in tribe Heliantheae. Our phylogenetic analysis of chloroplast DNA ndhF sequence data, however, reveals that it is part of tribe Astereae. In order to assess the relationships of this genus within the tribe, we produced a phylogeny based on ITS (nrDNA) sequence data of a sample including Amellus, African Conyza, Chrysocoma, Felicia spp., Mairia, Poeciliopsis, Printzia, Welwitschiella and Zyrphelis. Both parsimony and Bayesian analyses were done. The Bayesian analysis showed that African genera form a basal grade in tribe Astereae along with the Chinese Nannoglottis and South American and New Zealand genera, with Printzia being the earliest diverging member of the tribe. Mairia occupies an isolated position. Amellus, Chrysocoma, Felicia, Poecilolepis and Zyrphelis belong to subtribe Homochrominae, a South African radiation that also includes the St Helena endemics Commidendron and Melanodendron. Pteronia appears isolated, though it might be close to the Homochrominae. Welwitschiella is placed in the latest diverging African clade, subtribe Grangeinae, which also includes Grangea, Psiadia, Nidorella, and the African Conyza species except C. gouani. This subtribe is sister to the Eurasiatic subtribe Bellidinae, and together they are sister to the Astereae crown lineages of Australasia-Asia and South and North America.     

Chromosome and breeding system evolution of the genus Mercurialis (Euphorbiaceae): implications of ITS molecular phylogeny

 The internal transcribed spacers (ITS1 and ITS2) of nuclear ribosomal DNA were amplified and sequenced from 19 samples representing all species of the genus Mercurialis and two outgroup species, Ricinus communis and Acalypha hispida. The length of ITS1 in the ingroups ranged from 223 to 246 bp and ITS2 from 210 to 218 bp. Sequence divergence between pairs of species ranged from 1.15% to 25.88% among the ingroup species in the combined data of ITS1 and ITS2. Heuristic phylogenetic analyses using Fitch parsimony on the combined data of ITS1 and ITS2 with gaps treated as missing generated 45 equally parsimonious trees. The strict consensus tree was principally concordant with morphological classification. Within the genus, the ITS sequences recognised two main infrageneric clades: the M. perennis complex including three Eurasian stoloniferous species (M.␣leiocarpa, M. ovata and M. perennis) and the western Mediterranean group including eight both annual and perennial species. Of the western Mediterranean clade, the annual and perennial species grouped respectively into two different groups, and the annual life form is revealed as a synapomorphic character derived from perennial, whereas in the Eurasian clade ITS phylogeny suggested M. leiocarpa as basal clade sister to M.␣perennis and M. ovata. ITS phylogeny failed to resolve the relationships among the different cytotypes of M. ovata and M. perennis. ITS phylogeny also suggested rapid karyotypic evolution for the genus. The karyotypic divergence among the perennial species of western Mediterranean region did not corroborate the nucleotide sequence divergence among the species. Optimisation of chromosome numbers onto the ITS phylogeny suggested x=8 to be the ancestral basic chromosome number of the genus. ITS phylogeny confirmed that the androdioecy of M. ambigua is derived from dioecy. The nucleotide heterozygosity and additivity in ITS sequences clearly confirm the interspecific hybridisation in the genus Mercurialis. Received December 22, 2001; accepted May 21, 2002?Published online: November 14, 2002 Address of the authors: Martin Kr?henbühl, Yong-Ming Yuan (correspondence) and Philippe Küpfer, Institut de Botanique, Laboratoire de botanique évolutive, Université de Neuchatel, Emile-Argand 11, CH-2007 Neuchatel, Suisse. (e-mail: yong-ming.yuan@unine.ch)     

Piecing together the "new" Plantaginaceae

Scrophulariaceae is one of the families that has been divided extensively due to the results of DNA sequence studies. One of its segregates is a vastly enlarged Plantaginaceae. In a phylogenetic study of 47 members of Plantaginaceae and seven outgroups based on 3561 aligned characters from four DNA regions (the nuclear ribosomal ITS region and the plastid trnL-F, rps16 intron, and matK-trnK intron regions), the relationships within this clade were analyzed. The results from parsimony and Bayesian analyses support the removal of the Lindernieae from Gratioleae to a position outside Plantaginaceae. A group of mainly New World genera is paraphyletic with respect to a clade of Old World genera. Among the New World taxa, those offering oil as a pollinator reward cluster together. Ourisia is sister to this clade. Gratioleae consist of Gratiola, Otacanthus, Bacopa, Stemodia, Scoparia, and Mecardonia. Cheloneae plus Russelia and Tetranema together constitute the sister group to a clade predominantly composed of Old World taxa. Among the Old World clade, Ellisiophyllum and Lafuentea have been analyzed for the first time in a molecular phylogenetic analysis. The former genus is sister to Sibthorpia and the latter is surprisingly the sister to Antirrhineae.     

Phylogenetic reticulation in subtribe Helianthinae

Incongruence between phylogenetic estimates based on nuclear and chloroplast DNA (cpDNA) markers was used to infer that there have been at least two instances of chloroplast transfer, presumably through wide hybridization, in subtribe Helianthinae. One instance involved Simsia dombeyana, which exhibited a cpDNA restriction site phenotype that was markedly divergent from all of the other species of the genus that were surveyed but that matched the restriction site pattern previously reported for South American species of Viguiera. In contrast, analysis of sequence data from the nuclear ribosomal DNA internal transcribed spacer (ITS) region showed Simsia to be entirely monophyletic and placed samples of S. dombeyana as the sister group to the relatively derived S. foetida, a result concordant with morphological information. A sample of a South American species of Viguiera was placed by ITS sequence data as the sister group to a member of V. subg. Amphilepis, which was consistent with cpDNA restriction site data. Samples of Tithonia formed a single monophyletic clade based on ITS sequence data, whereas they were split between two divergent clades based on cpDNA restriction site analysis. The results suggested that cpDNA transfer has occurred between taxa diverged to the level of morphologically distinct genera, and highlight the need for careful and complete assessment of molecular data as a source of phylogenetic information.     

The phylogenetic relationships and evolution of the Canarian laurel forest endemicIxanthus viscosus (Aiton) Griseb. (Gentianaceae): Evidence frommatK and ITS sequences, and floral morphology and anatomy

The phylogenetic relationships of the Canarian laurel forest endemicIxanthus viscosus (Aiton) Griseb. (Gentianaceae) are investigated through a cladistic analysis of sequence variation of parts of the chloroplast genematK and the ITS region of nuclear ribosomal DNA. The floral anatomical and morphological characteristics ofIxanthus based on paraffin sections and SEM studies are also discussed. In the molecular analysesIxanthus is part of a clade of mostly temperate Erythraeinae and Chironiinae sensu Gilg, in which it is sister to the mostly Mediterranean annualBlackstonia. This relationship is supported by farreaching similarities in flower morphology and anatomy, and the shared possession of at least some perfoliate leaves. This finding contradicts most former classifications in whichIxanthus was placed either in subtribe Gentianinae which never includedBlackstonia, or in a subtribe of its own which was suggested to connect woody tropical with herbaceous temperate taxa. The phylogenetic relationships found here show that 1) the sister group ofIxanthus is distributed in the Mediterranean area, and that 2) its habit as a basally woody suffrutescent herb is best interpreted as an advanced character state.     

Molecular phylogenetics of tribe Synandreae, a North American lineage of lamioid mints (Lamiaceae)

The five mint genera Brazoria, Macbridea, Physostegia, Synandra and Warnockia (Lamioideae: Lamiaceae) are all North American endemics. Together with the monotypic European genus Melittis and the Asian genus Chelonopsis, these taxa have been classified as subtribe Melittidinae. Previous morphological studies have failed to uncover synapomorphic characters for this group. We sequenced the plastid trnL‐trnF region and trnS‐trnG spacer and the nuclear ribosomal 5S non‐transcribed spacer (5S‐NTS) to assess phylogenetic relationships within Melittidinae. Standard parsimony and direct optimization (POY) analyses show Melittis, the type genus of the subtribe, as sister to Stachys. Thus, the monophyly of subtribe Melittidinae is not supported either by molecular or morphological data. However, the North American endemics form a monophyletic group that can be recognized as the recircumscribed tribe Synandreae. The molecular relationships among these genera are corroborated by both morphological and cytological data. The expected close relationship between the south‐central endemics Warnockia and Brazoria and their sister relationship to the widespread genus Physostegia is confirmed. Nevertheless, most of the North American endemics are restricted to the south‐east of the continent. Dispersal westwards and northwards is correlated with an increase in chromosome numbers. No specific Eurasian origin (i.e., transatlantic or transpacific) can be determined, but Synandreae are clearly distinct from the large Stachys clade, and therefore represent a separate migration into North America. © The Willi Hennig Society 2007.     

Phylogenetic Analysis and Karyotype Evolution in the Genus Clivia(Amaryllidaceae)

Phylogenetic relationships of five taxa of Clivia, one probablenew species plus four recognized species, and three outgroupspecies were studied using sequences of the nuclear ribosomal5S non-transcribed spacer and the internal transcribed spacer(ITS) of 45S rDNA. Analysis of the data sets separately generatedsome well-supported groupings and congruent phylogenies. Cliviaminiata and C. gardenii are closely related. ‘Robust Gardenii’,the putative new species, is a sister clade of this group. Clivianobilis is distantly related to these three taxa and C. caulescensoccupies an intermediate position between the two groups. Chromosomelocations and distribution patterns of the 5S nuclear ribosomalgene in the species of Clivia were investigated using fluorescenceinsitu hybridization (FISH). In all species, only one pair of5S rDNA signals was observed. These were located on the shortarm of chromosome 8, at the position of the interstitial C-bands.The phylogenies obtained from the DNA sequences together withthe chromosome data accumulated here and previously publishedinformation on the location of the 45S rDNA sites have beenused to postulate evolutionary trends in Clivia chromosomes.Copyright 2001 Annals of Botany Company Clivia, chromosome evolution, 45S and 5S rDNA, ITS, FISH, molecular phylogeny     

The Bryophyta (Mosses): Systematic and Evolutionary Inferences from an rps4 Gene (cpDNA) Phylogeny

Phylogenetic analyses of nucleotide and amino acid sequencesof the chloroplast protein coding generps4 were performed for225 species of mosses, representing 84% of families recognizedby Vitt (1984. In: Schuster RM, ed. New manual of bryology,vol 2. Nichinan: Hattori Botanical Laboratory), under the criterionof maximum parsimony with Takakia and Sphagnum as outgroups.Most parsimonious topologies converge to a scenario whereinthe Andreaeidae are monophyletic and sister to the Bryidae (peristomatemosses), the Nematodonteae and the Buxbaumiaceae form a monophyleticlineage, the Diphysciaceae are sister to the Arthrodonteae and,within the latter, the Funarineae-Encalyptineae-Timmiaceae-Haplolepideaecompose a monophyletic clade sister to remaining diplolepideousmosses. This hypothesis suggests that early in the evolutionof the Arthrodonteae, two major lineages diverged, with oppositeand alternate peristomes, respectively. Bootstrap support forthe deep dichotomies is poor or lacking but increases when proteintranslations ofrps 4 sequences are included in the analysis.Several novel systematic hypotheses are raised, including (a)a diplolepideous rather than haplolepideous origin of the Pleurophascaceae;(b) an affinity of the Catascopiaceae with the Funariineae ratherthan the Bryineae; and (c) a close relationship of the Calomniaceaeand Mitteniaceae to the Rhizgoniaceae. The advantages and disadvantagesof a single gene phylogeny are discussed with respect to theidentification of polyphyletic familial or suprafamilial taxa.Copyright 2001 Annals of Botany Company Bryophyta, mosses, phylogeny, rps4, evolution, peristome, parsimony, sequences     

Polyphyly of the grass tribe Hainardieae (Poaceae: Pooideae): identification of its different lineages based on molecular phylogenetics, including morphological and cytogenetic characteristics

The small pooid grass tribe Hainardieae comprises six genera with approximately ten species; however, this tribe was not accepted by all previous taxonomic treatments. To study the relationships among these genera and to infer the phylogeny and evolutionary patterns, we used sequence variation of the internal transcribed spacers (ITS) of nuclear ribosomal and chloroplast (cp) matK DNA and morphology. Many genera of the Aveneae/Poeae tribe complex additionally were included. Both molecular datasets showed Hainardieae to be highly polyphyletic, and its genera to branch with different groups of the Aveneae/Poeae. Parapholis and Hainardia are corroborated as being closely related, and belonging to a firmly supported Eurasian clade together with Catapodium incl. Scleropoa, Cutandia, Desmazeria, Sphenopus, Vulpiella (subtribe Parapholiinae) and with Cynosurus as sister to this assemblage. The other genera of traditionally recognised Hainardieae are positioned phylogenetically distant: Mediterranean Narduroides is verified as more or less related to Festuca and relatives (subtribe Loliinae), whereas the west Eurasian Pholiurus is close to the lineage of Poa and relatives (subtribe Poinae). North American Scribneria is sister to Deschampsia and both genera should be unified under a common subtribe (Aristaveninae or Holcinae). The phylogenetic position of the Algerian genus Agropyropsis (close to Narduroides and within the Loliinae) is suggested on morphology only, because no molecular data was obtained for it. Considering classification, we support the abandonment of tribe Hainardieae and argue to abandon Poeae subtribe Scribneriinae. Poeae subtribe Parapholiinae is redefined with a novel genus content, due to the exclusion of Agropyropsis and Pholiurus and the inclusion of Vulpiella.     

Phylogenetic analysis of the subfamily Alpinioideae (Zingiberaceae), particularly Etlingera Giseke, based on nuclear and plastid DNA

Parsimony analyses based on DNA sequence data of the plastid group II intron rps16 and the internal transcribed spacer (ITS) were performed in order to examine the relationship of the pantropical subfamily Alpinioideae in Zingiberaceae (Zingiberales). Special emphasis was given to the large genus Etlingera placed in the tribe Alpinieae. A total of 50 taxa were included in the analysis. The strict consensus tree obtained by combining all data (280 parsimony informative characters of ITS, rps16, and coded indels) is well resolved with strongly supported clades. The subfamily Alpinioideae (excluding Pommereschea and Rhynchanthus) is strongly supported as monophyletic. The basal part of the tree is unresolved but a clade containing the derived genera of Alpinieae (Geocharis, Amomum, Hornstedtia, and Etlingera) is strongly supported. The establishment of Etlingera as the inclusive name for Achasma, Geanthus, and Nicolaia is also strongly supported: Etlingera is monophyletic with Hornstedtia as sister group.     

Phylogenetic relationships among Strobilanthes s.l. (Acanthaceae): evidence from ITS nrDNA, trnL-F cpDNA, and morphology

Chloroplast trnL-F sequence data, nuclear ribosomal internal transcribed spacer (ITS) sequence data, and morphology were used to analyze phylogenetic relationships among members of the subtribe Strobilanthinae. Parsimony and maximum likelihood analyses of trnL-F indicate that the Strobilanthinae are a monophyletic group. While parsimony analysis of ITS recovers a nonmonophyletic subtribe, maximum likelihood analysis of ITS corroborates results from trnL-F and suggests that systematic error is impacting on ITS parsimony analysis. A combined ITS and trnL-F analysis strengthens the signal and also recovers a monophyletic subtribe. All analyses indicate that Hemigraphis, Sericocalyx, and Strobilanthes are nonmonophyletic. With one exception, all morphological characters included in a combined ITS and morphological analysis are homoplastic. The prospect for a new informative generic classification of the Strobilanthinae aiming to recognize and diagnose only monophyletic groups is considered. While some groups can be diagnosed, adequate diagnosis of the majority of groups remains problematic. Consequently, a single expanded genus Strobilanthes sensu lato is proposed at the level of the well-supported and monophyletic Strobilanthinae.     

A MOLECULAR PHYLOGENY OF NORTH AMERICAN PLEUROCERIDAE (GASTROPODA: CERITHIOIDEA) BASED ON MITOCHONDRIAL 16S rDNA SEQUENCES

The Pleuroceridae Fischer, 1885, is one of three freshwatergastropod families currently recognized in the superfamily CerithioideaFérussac, 1819 (Mollusca: Caenogastropoda Cox, 1960).Despite considerable literature justifying various proposedgeneric names of North American pleurocerids, no study has beenconducted examining phylogenetic relationships of the recognizedgenera. In an effort to expand our understanding of evolutionaryrelationships of North American pleurocerid genrea, we examineda large portion of the mitochondrial 16S rRNA gene among 32extant North American taxa. Multiple sequence alignment of theamplified region for our taxa resulted in a matrix consistingof 900 nucleotides including insertions and deletions. Basedon analysis of nucleotide substitution patterns, we employedtwo approaches in our phylogenetic analysis: (1) all substitutionsreceived equal weighting and (2) transversions were weighted2X and 4X transitions to compensate for transition saturationamong distantly related taxa. The molecular phylogeny basedon the mitochondrial 16S rDNA sequences supports the monophylyof Pleurocera Rafinesque, 1819, Elimia H. & A. Adams, 1854,and Juga H. & A. Adams, 1854, but depicts the genera LithasiaHaldeman, 1840, and Leptoxis Rafinesque, 1819, as polyphyletic.The genus Pleurocera is sister to Elimia, which in turn is sisterto a paraphyletic assemblage including representatives of Leptoxis,Lithasia, and the monotypic genus Io Lea, 1831. Juga, a genusrestricted to west of the North American continental divideis the basal-most clade and is sister to all the aforementionedgenera found east of the continental divide. (Received 26 January 1999; accepted 14 October 1999)     

Systematics of North American Froelichia (Amaranthaceae subfam. Gomphrenoideae) II: Phylogeny and biogeographic speciation patterns inferred from nrITS sequence data

Molecular phylogenetic analyses of the nuclear ribosomal DNA internal transcribed spacer (ITS 1 and ITS 2) and the 5.8S gene were used to infer a phylogeny among the ten recognized taxa of Froelichia in North America. Analyses using both maximum parsimony (MP) and maximum-likelihood (ML) depicted a low level of sequence divergence though it was sufficient in most cases to differentiate taxa. Froelichia xantusii, a species restricted to southern Baja California was shown to be the basalmost member of the group subtending three clades. Two of the clades received good bootstrap support in the MP analysis and corresponded to a genetically homogeneous F. interrupta, and a clade comprising the two species F. latifolia and F. texana. A third clade receiving low bootstrap support contained F. floridana, F. gracilis, F. arizonica, and F. drummondii. Species diversity within the genus was centered within the Tamaulipan Brushland region of north-east Mexico and the southern portion of the US state of Texas where taxa from two of the three principal clades occurred, indicating a region of high speciation and diversification within the genus.     

a molecular phylogeny of apiaceae subfamily Apioideae: evidence from nuclear ribosomal DNA internal transcribed spacer sequences

Phylogenetic relationships among 40 New World and Old World members of Apiaceae subfamily Apioideae, representing seven of the eight tribes and eight of the ten subtribes commonly recognized in the subfamily, were inferred from nucleotide sequence variation in the internal transcribed spacer (ITS) regions of 18-26S nuclear ribosomal DNA. Although the sequences are alignable, with only 11% of sites excluded from the analyses because of alignment ambiguity, divergence values in pairwise comparisons of unambiguous positions among all taxa were high and ranged from 0.5 to 33.2% of nucleotides in ITS 1 and from 0 to 33.2% of nucleotides in ITS 2. Average sequence divergence across both spacer regions was 18.4% of nucleotides. Phylogenies derived from ITS sequences estimated using neighbor-joining analysis of substitution rates, and maximum likelihood and parsimony methods give trees of essentially similar topology and indicate that: (1) there is little support for any existing system of classification of the subfamily that is based largely on morphological and anatomical features of the mericarp; (2) there is a major phylogenetic division within the subfamily, with one clade comprising the genus Smyrnium and those taxa belonging to Drude's tribes Dauceae, Scandiceae, and Laserpitieae and the other clade comprising all other examined taxa; and (3) the genera Arracacia, Coaxana, Coulterophytum, Enantiophylla, Myrrhidendron, Prionosciadium, and Rhodosciadium, all endemic to Mexico and Central America, comprise a clade but their relationships to other New World taxa are equivocal. A phylogeny derived from parsimony analysis of chloroplast DNA rpoC1 intron sequences is consistent with, but considerably less resolved than, relationships derived from these ITS regions. This study affirms that ITS sequences are useful for phylogenetic inference among closely related members of Apioideae but, owing to high rates of nucleotide substitution, are less useful in resolving relationships among the more ancestral nodes of the phylogeny.     

Origin and evolution of the endemic Macaronesian Inuleae (Asteraceae): evidence from the internal transcribed spacers of nuclear ribosomal DNA

The tribe Inuleae (Asteraceae) has 10 species endemic to the Macaronesian islands, including the three endemic genera Allagopappus, Schizogyne, and Vierea. Phylogenetic analyses of DNA sequence data from the internal transcribed spacers (ITS) of the nuclear ribosomal DNA of 47 taxa were performed using all Macaronesian endemics and representative species from 21 of the 36 genera of the Inuleae. The resulting ITS phylogeny reveals that Allagopappus is sister to a large clade that contains all genera with a predominantly Mediterranean distribution. This finding suggests that Allagopappus may represent an ancient lineage that found refuge in the Canary Islands following the major climatic and/or geologic changes in the Mediterranean basin after the Tertiary. The Macaronesian endemic genus Schizogyne is sister to Limbarda from the Mediterranean. The third Macaronesian endemic genus, Vierea, is sister to Perralderia, which is restricted to Morocco and Algeria. Pulicaria canariensis is sister to P. mauritanica, a species endemic to Morocco and Algeria. In contrast, P. diffusa from the Cape Verde Islands is sister to a broadly distributed species, P. crispa, that occurs from North Africa to the Arabian peninsula. Based on the ITS data, the genera Blumea, Inula, and Pulicaria are not monophyletic. The ITS trees suggested that Blumea mollis belongs to the tribe Plucheeae, a finding that is congruent with recent morphological evidence. A possible southern African origin for the core of the Laurasian taxa of the Inuleae is also suggested.     

Evidence for host-specific clades of tetraphyllidean tapeworms (Platyhelminthes: Eucestoda) revealed by analysis of 18S ssrDNA

Sequence data from the V4 and V7-V9 variable regions of the 18S small subunit ribosomal DNA (ssrDNA) gene were used to examine relationships among 26 tetraphyllidean and two lecanicephalidean taxa. Newly collected specimens of 21 of the tetraphyllidean species were used to generate ssrDNA sequences that were combined with sequences previously available, including those of two diphyllidean taxa used for outgroup rooting. The sequences were aligned by eye according to secondary structural motifs of the conserved core of the molecule. Of the 1520 sites in the alignment, 874 (58%) were excluded from analysis due to alignment gaps and lack of positional homology as inferred by manual inspection. Genetic variability of the ssrDNA gene regions compared was greater than would be expected, based on the present taxonomy of the ingroup species, and the genetic divergences among tetraphyllidean 'families' and genera were comparable to that among tapeworm orders. Phylogenetic hypotheses were generated by the methods of maximum parsimony and maximum likelihood (GTR + I + Gamma nucleotide substitution model). Four most parsimonious trees resulted from analysis by maximum parsimony. Strict consensus of the four trees supported the monophyly of the Tetraphyllidea, with the lecanicephalidean taxa forming a sister lineage. Among the tetraphyllidean taxa included in the analysis were three major clades: a basal clade including species of the phyllobothriid genera Anthocephalum, Echeneibothrium, Rhinebothrium, Rhodobothrium and Spongiobothrium; a clade uniting the phyllobothriids of the genus Duplicibothrium with the dioecotaeniid genus Dioecotaenia; and a larger sister clade to the Duplicibothrium + Dioecotaenia clade that included the phyllobothriid genera Caulohothrium, Ceratobothrium, Clistobothrium, Paraoryigmatobothrium and Prosobothrium, the litobothriid genus Litobothrium and the onchobothriid genera Acanthobothrium, Calliobothrium, Phoreiobothrium and Platybothrium. Maximum likelihood analysis resulted in a topology that was congruent where nodes were strongly supported by parsimony analysis, but differed in the relative positions of the well-supported clades. In addition,maximum likelihood analysis grouped the lecanicephalidean taxa among the tetraphyllidean taxa, indicating paraphyly of the order Tetraphyllidea as currently defined. Relationships suggested by both methods of analysis reflected common host associations of the taxa better than their current classification, suggesting that coevolution has had a significant role in the evolution of the group.     

A molecular phylogenetic study of the subtribe Glycininae (Leguminosae) derived from the chloroplast DNA rps16 intron sequences

Phylogenetic relationships among 13 genera of the subtribe Glycininae, two genera of the allied subtribe Diocleinae that were included within Glycininae by Polhill, and two genera of the subtribe Erythrininae as outgroups were inferred from chloroplast DNA rps16 intron sequence variation. Pairwise sequence divergence values ranged from identity between Teramnus mollis and T. micans and between T. flexilis and T. labialis to 7.89% between Pueraria wallichii and Pseudeminia comosa across all accessions. Phylogenies estimated using parsimony and neighbor-joining methods revealed that (1) Glycininae is monophyletic if Pachyrhizus and Calopogonium (both Diocleinae) are included within Glycininae; (2) the genus Teramnus is closely related to Glycine, and Amphicarpaea showed a sister relationship to the clade comprising Teramnus and Glycine; (3) the expanded Glycininae including two genera of Diocleinae is divided into three branches, temporarily named I (comprising the rest of the examined taxa), II (Pueraria wallichii), and III (Mastersia), but their relationships are equivocal; and (4) the genus Pueraria, regarded as a closely related genus to Glycine, is not monophyletic and should be divided into at least four genera (a hypothesis supported previously by Lackey).