Phylogenetic and biogeographic diversification of Rhus (Anacardiaceae) in the Northern Hemisphere

Sequences of internal transcribed spacers (ITS) of nuclear ribosomal DNA, the chloroplast ndhF gene, and chloroplast trnL-F regions (trnL intron, and trnL [UAA] 3' exon-trnF [GAA] intergenic spacer) were used for phylogenetic analyses of Rhus, a genus disjunctly distributed in Asia, Europe, Hawaii, North America, and Northern Central America. Both ITS and cpDNA data sets support the monophyly of Rhus. The monophyly of subgenus Rhus was suggested by the combined cpDNA and ITS data, and largely supported in the cpDNA data except that Rhus microphylla of subgenus Lobadium was nested within it. The monophyly of subgenus Lobadium was strongly supported in the ITS data, whereas the cpDNA data revealed two main clades within the subgenus, which formed a trichotomy with the clade of subgenus Rhus plus R. microphylla. The ITS and cpDNA trees differ in the positions of Rhus michauxii, R. microphylla, and Rhus rubifolia, and hybridization may have caused this discordance. Fossil evidence indicates that Rhus dates back to the early Eocene. The penalized likelihood method was used to estimate divergence times, with fossils of Rhus subgenus Lobadium, Pistacia and Toxicodendron used for age constraints. Rhus diverged from its closest relative at 49.1+/-2.1 million years ago (Ma), the split of subgenus Lobadium and subgenus Rhus was at 38.1+/-3.0 Ma. Rhus most likely migrated from North America into Asia via the Bering Land Bridge during the Late Eocene (33.8+/-3.1 Ma). Rhus coriaria from southern Europe and western Asia diverged from its relatives in eastern Asia at 24.4+/-3.2 Ma. The Hawaiian Rhus sandwicensis diverged from the Asian Rhus chinensis at 13.5+/-3.0 Ma. Subgenus Lobadium was inferred to be of North American origin. Taxa of subgenus Lobadium then migrated southward to Central America. Furthermore, we herein make the following three nomenclatural combinations: (1) Searsia leptodictya (Diels) T. S. Yi, A. J. Miller and J. Wen, comb. nov., (2) Searsia pyroides (A. Rich.) T. S. Yi, A. J. Miller and J. Wen, comb. nov., and (3) Searsia undulata (Jacq.) T. S. Yi, A. J. Miller and J. Wen, because our analyses support the segregation of Searsia from Rhus.     

Chloroplast-DNA restriction site analysis in the genus Bromus (Poaceae)

Chloroplast DNA (cpDNA) restriction site variation was examined in 32 species, representing five subgenera, of Bromus (Poaceae). Thirty-seven phylogenetically informative restriction sites were detected. Cladistic analysis of the restriction site data produced a single most-parsimonious tree of 50 steps. The cladogram indicated two major clades within the genus. One clade included B. trinii of subgenus Neobromus and species of subgenus Ceratochloa. The other was composed of subgenera Festucaria, Stenobromus, and Bromus. Within the second clade, species of subgenus Festucaria appeared in three lineages. The second clade also contained an assemblage of species belonging to subgenera Stenobromus and Bromus in a separate lineage. There was very little resolution of relationships in this assemblage since several species appeared individually in separate lineages. The cpDNA phylogenetic hypothesis did not separate species of subgenera Stenobromus and Bromus into well-defined clades as circumscribed by morphology and cytogenetics. The cpDNA tree is in agreement with the phylogenetic scheme based on traditional data in that: 1) subgenera Neobromus and Ceratochloa were the first to diverge, while Bromus and Stenobromus diverged later; 2) within the genus Bromus species with small chromosomes are ancestral; and 3) subgenera Bromus and Stenobromus probably originated from similar ancestors as Festucaria. The tree based on cpDNA data does not support that: 1) subgenera Neobromus and Ceratochloa did not have a common origin; 2) subgenus Festucaria is monophyletic; and 3) subgenera Stenobromus and Bromus are distinct entities. The mean nucleotide sequence divergence values between pairs of subgenera ranged from p = 0.0 to 0.9. These values suggest that cpDNA evolution in Bromus is slow.     

Phylogeny and infrageneric classification of Symplocos (Symplocaceae) inferred from DNA sequence data

Symplocos comprises ～300 species of woody flowering plants with a disjunct distribution between the warm-temperate to tropical regions of eastern Asia and the Americas. Phylogenetic analyses of 111 species of Symplocos based on the nuclear ribosomal internal transcribed spacer (ITS) region and the chloroplast genes rpl16, matK, and trnL-trnF yielded topologies in which only one of the four traditionally recognized subgenera (Epigenia; Neotropics) is monophyletic. Section Cordyloblaste (subgenus Symplocos; eastern Asia) is monophyletic and sister to a group comprising all other samples of Symplocos. Section Palura (subgenus Hopea; eastern Asia) is sister to a group comprising all other samples of Symplocos except those of section Cordyloblaste. Symplocos wikstroemiifolia (eastern Asia) and S. tinctoria (southeastern United States), both of subgenus Hopea, form a clade that groups with S. longipes (tropical North America) and the species of subgenus Epigenia. The remaining samples of subgenus Hopea (eastern Asia) form a clade. Section Neosymplocos (subgenus Microsymplocos; Neotropics) is well nested within a clade otherwise comprising the samples of section Symplocastrum (subgenus Symplocos; Neotropics). Section Urbaniocharis (subgenus Microsymplocos; Antilles) groups as sister to the clade comprising Symplocastrum and Neosymplocos. The data support the independent evolution of deciduousness among section Palura and S. tinctoria. The early initial divergence of sections Cordyloblaste and Palura from the main group warrants their recognition at taxonomic levels higher than those at which they are currently placed. An inferred eastern Asian origin for Symplocos with subsequent dispersal to the Americas is consistent with patterns from other phylogenetic studies of eastern Asian-American disjunct plant groups but contrary to a North American origin inferred from the earliest fossil occurrences of the genus.     

A CHLOROPLAST-DNA PHYLOGENY OF THE WILD PERENNIAL RELATIVES OF SOYBEAN (GLYCINE SUBGENUS GLYCINE): CONGRUENCE WITH MORPHOLOGICAL AND CROSSING GROUPS

Hypotheses of evolutionary relationships among the Australian wild perennial relatives of soybean (Glycine subgenus Glycine) are based largely on patterns of meiotic pairing in intra- and interspecific experimental hybrids. This evidence has indicated a number of genome groupings within the subgenus but has not resolved most phylogenetic relationships. Restriction-endonuclease site variation of chloroplast DNA (cpDNA) within the perennial subgenus is reported here, representing a sampling of approximately 3% of the approximately 150-kilobase plastome. Seven hundred twenty-one unique restriction sites were compared within Glycine using 29 restriction endonucleases; 157 sites varied within the genus. Distance and parsimony methods using these data yielded congruent results, recognizing the existence of three major groups within subgenus Glycine: the species-rich and geographically diverse A clade consisting of G. canescens and related taxa; the B clade, which includes the stoloniferous species; and the C group, containing two species with distinctive curved pods. These results are in general agreement with hypotheses based on genome analysis; inconsistencies involve the inclusion of genetically divergent taxa such as G. falcata in well-supported plastome clades comprised of otherwise interfertile species. Such findings are not unexpected if crossing barriers are considered to be unique features of such anomalous species, paralleling their often numerous morphological and cpDNA autapomorphies. Consideration of cpDNA divergence within the three major clades of subgenus Glycine indicates that the rate of plastome evolution is uncoupled from rates of morphological or ecological diversification.     

Differential organellar inheritance in Passiflora’s (Passifloraceae) subgenera

Four chloroplast (cp), one mitochondrial (mt), and one ribosomal nuclear (ITS) DNA regions were studied in four artificial and one natural interspecific Passiflora hybrids. The ITS results confirmed their hybrid origin and all mtDNAs were maternally inherited. The same, however, was not true for cpDNA. The four hybrids (three artificial and one natural) derived from species of the Passiflora subgenus showed a cpDNA paternal inheritance, while the one involving taxa of the Decaloba subgenus gave evidence of maternal transmission. These results are of significance for the ongoing studies which are being performed on the molecular evolution of this genus and furnish important background for investigations aimed at clarifying the factors which determine cpDNA inheritance.     

Molecular phylogenetics and evolution of the endemic Hawaiian genus Adenophorus (Grammitidaceae)

Recent studies of the phylogeny of several groups of native Hawaiian vascular plants have led to significant insights into the origin and evolution of important elements of the Hawaiian flora. No groups of Hawaiian pteridophytes have been subjected previously to rigorous phylogenetic analysis. We conducted a molecular phylogenetic analysis of the endemic Hawaiian fern genus Adenophorus employing DNA sequence variation from three cpDNA fragments: rbcL, atpbeta, and the trnL-trnF intergenic spacer (IGS). In the phylogenetic analyses we employed maximum parsimony and Bayesian inference. Bayesian phylogenetic inference often provided stronger support for hypothetical relationships than did nonparametric bootstrap analyses. Although phylogenetic analyses of individual DNA fragments resulted in different patterns of relationships among species and varying levels of support for various clades, a combined analysis of all three sets of sequences produced one, strongly supported phylogenetic hypothesis. The primary features of that hypothesis are: (1) Adenophorus is monophyletic; (2) subgenus Oligadenus is paraphyletic; (3) the enigmatic endemic Hawaiian species Grammitis tenella is strongly supported as the sister taxon to Adenophorus; (4) highly divided leaf blades are evolutionarily derived in the group and simple leaves are ancestral; and, (5) the biogeographical origin of the common ancestor of the Adenophorus-G. tenella clade remains unresolved, although a neotropical origin seems most likely.     

Molecular phylogenetics of the clover genus (Trifolium--Leguminosae)

Trifolium, the clover genus, is one of the largest genera of the legume family. We conducted parsimony and Bayesian phylogenetic analyses based on nuclear ribosomal DNA internal transcribed spacer and chloroplast trnL intron sequences obtained from 218 of the ca. 255 species of Trifolium, representatives from 11 genera of the vicioid clade, and an outgroup Lotus. We confirm the monophyly of Trifolium, and propose a new infrageneric classification of the genus based on the phylogenetic results. Incongruence between the nrDNA and cpDNA results suggests five to six cases of apparent hybrid speciation, and identifies the putative progenitors of the allopolyploids T. dubium, a widespread weed, and T. repens, the most commonly cultivated clover species. Character state reconstructions confirm 2n=16 as the ancestral chromosome number in Trifolium, and infer a minimum of 19 instances of aneuploidy and 22 of polyploidy in the genus. The ancestral life history is hypothesized to be annual in subgenus Chronosemium and equivocal in subgenus Trifolium. Transitions between the annual and perennial habit are common. Our results are consistent with a Mediterranean origin of the genus, probably in the Early Miocene. A single origin of all North and South American species is hypothesized, while the species of sub-Saharan Africa may originate from three separate dispersal events.     

Are any primroses (Primula) primitively monomorphic?

Primula (c. 430 species) and relatives (Primulaceae) are paradigmatic to our understanding of distyly. However, the common co-occurrence of distyly and monomorphy in closely related groups within the family has made the interpretation of its evolution difficult.Here, we infer a chloroplast DNA (cpDNA) phylogeny for 207 accessions, including 51% of the species and 95% of the sections of Primula with monomorphic populations, using Bayesian methods. With this tree, we infer the distribution of ancestral states on critical nodes using parsimony and likelihood methods.The inferred cpDNA phylogeny is consistent with prior estimates. The most recent common ancestor (MRCA) of Primula is resolved as distylous using both methods of inference. However, whether the distyly in Primula, Hottonia, and Vitaliana arose once or three independent times is not clear.We conclude that monomorphism in descendants of the MRCA of Primula is derived from distyly in all cases. Thus, scenarios for the evolution of distyly that rely on the persistence of primitive monomorphy (such as in Primula section Sphondylia) require re-evaluation.     

不同类型水稻不育系药隔发育过程中Ca2+的分布变化

采用焦锑酸钾沉淀法研究了不同类型水稻不育系与其相应保持系及其可育花药药隔发育过程中Ca^2＋的分布变化。结果显示在正常花药的发育过程中,药隔中的Ca^2＋呈以下变化规律：（1）花药发育早期药隔中的Ca^2＋沉淀很少;（2）随着花药的发育,药隔中的Ca^2＋沉淀增加,木质部细胞的次生加厚壁上有较多的Ca^2＋沉淀,连接组织中的Ca^2＋沉淀也大大增加;（3）到了花粉内容物充实后期及成熟花粉时期,药隔中的Ca^2＋又略有减少。而不育花药药隔中的Ca^2＋在花药发育的各时期均比可育花药要多。败育类型不同,药隔中出现大量Ca^2＋沉淀的时期也不同,无花粉型不育系在花粉母细胞时期药隔中就有大量Ca^2＋沉淀,而农垦58S从单核花粉时期约隔中才出现大量Ca^2＋沉淀。实验结果表明：Ca^2＋在不同类型的不育水稻中呈现出一定的规律性变化,药隔中Ca^2＋的异常分布可能与水稻花粉的败育有关。     

Analyses of cpDNA matK sequence data place Tillaea (Crassulaceae) within Crassula

Analysis of cpDNA matK sequences for a total of 43 members of the succulent plant family Crassulaceae, including 24 taxa of Crassula, recovered a well-supported clade comprising Crassula species that is sister to the remainder of the family. The resulting topologies do not support the monophyly of the currently recognized subgenera of Crassula, as one member of subgenus Disporocarpa (C. crenulata) is placed as sister to an otherwise monophyletic subgenus Crassula. The major synapomorphy that has been used to recognize the latter subgenus is a base chromosome number of x = 7 versus a base of x = 8 in the other subgenus. We cannot assess the utility of this feature for defining subgenus Crassula because a chromosome count of C. crenulata has yet to be published. The five accessions of the recently resurrected segregate genus Tillaea (of 24 total Crassula species) included here were placed in four separate, well-supported lineages, one of which is greatly removed from the other four accessions. This suggests that this genus is not valid and should not be recognized. An initial examination of the evolution of habit indicates that a perennial habit is ancestral and that the annual habit is a feature that has been derived at least twice in the genus.     

Phylogeny and patterns of floral diversity in the genus Piper (Piperaceae)

With ～1000 species distributed pantropically, the genus Piper is one of the most diverse lineages among basal angiosperms. To rigorously address the evolution of Piper we use a phylogenetic analysis of sequences of the internal transcribed spacers (ITS) of nuclear ribosomal DNA based on a worldwide sample. Sequences from a total of 51 species of Piper were aligned to yield 257 phylogenetically informative sites. A single unrooted parsimony network suggested that taxa representing major geographic areas could potentially form three monophyletic groups: Asia, the South Pacific, and the Neotropics. The position of Pothomorphe was well supported among groups of New World taxa. Simultaneous phylogenetic analysis of an expanded alignment including outgroups suggested that taxa from the South Pacific and Asia formed a monophyletic group, provisionally supporting a single origin of dioecy. Within the Neotropical sister clade, resolution was high and strong bootstrap support confirmed the monophyly of several traditionally recognized infrageneric groups (e.g., Enckea [including Arctottonia], Ottonia, Radula, Macrostachys). In contrast, some of the species representing the highly polytypic subgroup Steffensia formed a clade corresponding to the previously recognized taxon Schilleria, while others were strongly associated with several of the more specialized groups of taxa. The distribution of putatively derived inflorescence and floral character states suggested that both umbellate and solitary axillary inflorescences have multiple origins. Reduction in anther number appears to be associated with highly packaged inflorescences or with larger anther primordia per flower, trends that are consistent with the suppression of later stages of androecial development.     

Molecular analysis of interspecific and intergeneric relationships of Banksia using RAPDs and non-coding chloroplast DNA sequences

 Random amplified polymorphic DNA (RAPD) and chloroplast DNA (cpDNA) markers were used to study interspecific and intergeneric relationships of Banksia (Proteaceae) to aid breeding of the genus for cut flower production. The accepted morphological phylogeny of Banksia, with two subgenera, two sections and 13 series, is unclear regarding the relationships of the commercial cut flower species B. coccinea. Fifteen RAPD primers and a non-coding cpDNA sequence between the trnL (UAA) and trnF (GAA) gene were applied to species of Banksia, the related genus Dryandra, and to Musgravea heterophylla as the outgroup, with cluster analysis applied to the results. The two methods were in broad agreement with each other, and with the accepted taxonomy, with closely related species pairs and groups clustering together, but RAPDs were not informative between distantly related species or species pairs. Banksia coccinea clustered with Dryandra and formed a polytomy with 2 Dryandra species and the two sections of subgenus Banksia. Subgenus Isostylis formed a polytomy with D. formosa, basal to subgenus Banksia, but with B. cuneata and B. illicifolia (both in subgenus Isostylis) polyphyletic. Dryandra did not separate as a clade and fell within Banksia, raising questions about the currently accepted view of the two as sister genera with parallel morphological development. The results indicate that interspecific and intergeneric hybridisation with genus Dryandra and subgenus Banksia may be possible routes for improvement of the commercial species B. coccinea. Received: 25 September 1996 / Accepted: 8 November 1996     

Incongruence between chloroplast and species phylogenies in Eucalyptus subgenus Monocalyptus (Myrtaceae)

Seventy-eight polymorphic cpDNA (chloroplast DNA) characters were found in 13 closely related taxa from Eucalyptus series Amygdalinae (subgenus Monocalyptus) and seven potential outgroup taxa. The strict consensus of six cladograms generated from cpDNA data confirmed monophyly of Monocalyptus. However, cpDNA phylogeny within Monocalyptus was incongruent with taxonomic classification, being more related to geography, even when accessions were from divergent series. Monocalyptus cpDNA formed two major clades. On the island of Tasmania cpDNA was restricted to a single clade, exhibited very little variation, and was phylogenetically related to cpDNA found in central and western Victoria. In contrast, cpDNA of mainland monocalypt taxa was more variable, even within the Amygdalinae. Four out of six Tasmanian Amygdalinae species were polymorphic. The difference between cpDNA of replicates was often greater than differences between species from different series. The low level of cpDNA variation and extensive morphological intergradation between the Tasmanian endemics suggest recent speciation. However, the transfer of cpDNA through hybridization between lineages is the most likely explanation for the observed sharing of cpDNA across series. This study highlights that the geographical pattern to cpDNA variation in Eucalyptus may be an important source of information on past plant distributions in Australia.     

A plastid DNA phylogeny of the genus Acacia Miller (Acacieae, Leguminoseae)

Past classifications of the tribe Acacieae Rchb. are outlined and the confusion concerning the relationships of the three subgenera of Acacia Mill. are highlighted. A plastid DNA analysis of Acacieae shows that the genus Acacia is not monophyletic. Furthermore subgenera Acacia Vassal and Aculeiferum Vassal are sister taxa and neither appear closely related to subgenus Phyllodineae (DC.) Ser. Subgenera Acacia and Aculeiferum form a clade that is basal to a well-supported clade consisting of tribe Ingeae Benth. taxa, Faidherbia albida (Del.) A. Chev. and subgenus Phyllodineae. The series of relationships suggested by the cpDNA data contradicts previous investigations of the tribe. Possible explanations of this conflict are explored, and the taxonomic implications of the plastid DNA data set are considered.     

Sequence variation of non-coding regions of chloroplast DNA of soybean and related wild species and its implications for the evolution of different chloroplast haplotypes

Soybean chloroplast DNAs (cpDNAs) are classified into three types (I, II and III) based on RFLP profiles. Type I is mainly observed in cultivated soybean (Glycine max), while type II and type III are frequently found in both cultivated and wild soybean (Glycine soja), although type III is predominant in wild soybean. In order to evaluate the diversity of cpDNA and to determine the phylogenetic relationship among different chloroplast types, we sequenced nine non-coding regions of cpDNA for seven cultivated and 12 wild soybean accessions with different cpDNA types. Eleven single-base substitutions and a deletion of five bases were detected in a total of 3849 bases identified. Five mutations distinguished the accessions with types I and II from those with type III, and seven were found in the accessions with type III, independently of their taxa. Four species of the subgenus Glycine shared bases that were identical to those with types I and II at two of the five mutation sites and shared bases that were identical to those with type III at the remaining three sites. Therefore, the different cpDNA types may not have originated monophyletically, but rather may have differentiated from a common ancestor in different evolutionary directions. A neighbor-joining tree resulting from the sequence data revealed that the subgenus Soja connected with Glycine microphylla which formed a distinct clade from Clycine clandestina and the tetraploid cytotypes of Glycine tabacina and Glycine tomentella. Several informative length mutations of 54 to 202 bases, due to insertions or deletions, were also detected among the species of the genus Glycine. Received: 16 December 1999 / Accepted: 12 February 2000     

Phylogenetic relationships among subgenera,species, and varieties of Japanese <Emphasis Type="Italic">Salvia</Emphasis> L. (Lamiaceae)

To determine evolutionary relationships among all Japanese members of the genus Salvia (Lamiaceae), we conducted molecular phylogenetic analyses of two chloroplast DNA (cpDNA) regions (rbcL and the intergenic spacer region of trnL−trnF:trnL−trnF) and one nuclear DNA (nrDNA) region (internal transcribed spacer, ITS). In cpDNA, nrDNA, and cpDNA+nrDNA trees, we found evidence that all Japanese and two Taiwanese Salvia species are included in a clade with other Asian Salvia, and Japanese Salvia species were distributed among three subclades: (1) S. plebeia (subgenus Sclarea), (2) species belonging to subg. Salvia, and (3) species belonging to subg. Allagospadonopsis. At the specific level our findings suggest: a close relationship between S. nipponica and S. glabrescens, no support for monophyly of S. lutescens and its varieties in cpDNA, nrDNA and cpDNA+nrDNA trees, and that S. pygmaea var. simplicior may be more closely related to S. japonica than to other varieties of S. pygmaea.     

Phylogeny, biogeography, and processes of molecular differentiation in Quercus subgenus Quercus (Fagaceae).

Quercus is one of the most abundant and economically important genera of woody plants in the Northern Hemisphere. To infer phylogenetic relationships within Quercus subgenus Quercus, chloroplast DNA (cpDNA) restriction sites and nucleotide sequences of the internal transcribed spacers (ITS) and the 5.8S coding region of the nuclear ribosomal DNA repeat were obtained for 44 individuals, including 25 species, intraspecific samples, and three outgroups. Separate parsimony analyses of each data set showed that individual gene trees were congruent and often complementary in supporting clades that generally corresponded to previously recognized taxonomic groups. Only one instance of strongly supported gene tree incongruence was detected and this anomalous pattern was explained best by ancient introgression of cpDNA across sectional boundaries. Simultaneous parsimony analysis of the pruned data sets supported the recognition of the strictly Eurasian section Cerris and resolved a novel hypothesis for the major infrageneric groups (Cerris- (Lobatae- (Protobalanus + Quercus sensu stricto))). The biogeographic hypothesis that all major oak lineages evolved locally at middle latitudes within the general distribution of their fossil ancestors was fully supported. This set of relationships also suggested a New World origin for the widespread white oaks of the Northern Hemisphere (section Quercus s. s.). For both data sets, inter- and intraspecific sampling within section Protobalanus showed little correspondence to morphological species. Greater cladistic structure among the samples was obtained by cpDNA restriction sites and two well-delimited plastomes types comprising a total of 15 distinct haplotypes were resolved. Haplotypes of 2 of the peripheral species in this species complex occupy terminal portions of one of the plastome clades, suggesting a more recent origin relative to those of more widespread species. The phylogeography of the two divergent plastome types suggested a north-south pattern, consistent with a Late Tertiary disjunction in the ancestral distribution of section Protobalanus.     

Phylogeny and biogeography of Rhododendron subsection Pontica, a group with a tertiary relict distribution

Rhododendron subgenus Hymenanthes subsection Pontica is exceptional among Tertiary relict groups in having a high proportion of species (4 of 11) native to SW Eurasia. A phylogeny based on cpDNA matK and trnL-F indicated that multiple Pontica lineages colonised each of SW Eurasia, SE North America, and NE Asia, with little or no speciation within regions thereafter. Therefore, multiple (3-4) Pontica lineages survived the Quaternary in SW Eurasia, in contrast to other Tertiary relict genera. Pontica comprises two major clades, one of which is wholly Eurasian, and paraphyletic with respect to at least some of the remaining 200 species of subgenus Hymenanthes, which are all distributed in SE Asia. The other clade has species from W and SE North America, SW Eurasia, and NE Asia. According to synonymous matK substitution data, the two clades diverged 9-6 million years ago (mya), whereas most divergence within them happened 5-3 mya. Although the phylogeny indicates probable trans-Atlantic migration for one of two America-Eurasia disjunctions in Pontica, the timing supports migration via Beringia for both.     

Origin and relationships of Saintpaulia (Gesneriaceae) based on ribosomal DNA internal transcribed spacer (ITS) sequences

Phylogenetic relationships of eight species of Saintpaulia H. Wendl., 19 species of Streptocarpus Lindl. (representing all major growth forms within the genus), and two outgroups (Haberlea rhodopensis Friv., Chirita spadiciformis W. T. Wang) were examined using comparative nucleotide sequences from the two internal transcribed spacers (ITS) of nuclear ribosomal DNA. The length of the ITS 1 region ranged from 228 to 249 base pairs (bp) and the ITS 2 region from 196 to 245 bp. Pairwise sequence divergence across both spacers for ingroup and outgroup species ranged from 0 to 29%. Streptocarpus is not monophyletic, and Saintpaulia is nested within Streptocarpus subgenus Streptocarpella. Streptocarpus subgenus Streptocarpus is monophyletic. The ITS sequence data demonstrate that the unifoliate Streptocarpus species form a clade, and are also characterized by a unique 47-bp deletion in ITS 2. The results strongly support the monophyly of (1) Saintpaulia, and (2) Saintpaulia plus the African members of the subgenus Streptocarpella of Streptocarpus. The data suggest the evolution of Saintpaulia from Streptocarpus subgenus Streptocarpella. The differences in flower and vegetative characters are probably due to ecological adaptation leading to a relatively rapid radiation of Saintpaulia.     

151 Phylogenetic Analysis of The Subgenus Euglena with Particualr Reference to the Type Species Euglena Viridis (Euglenophyceae)

Euglena viridis was first described by Antony van Leeuwenhoek in 1674. This taxon later became the type for the genus Euglena erected by Ehrenberg in 1838. The primary characters that distinguish this taxon are the single stellate chloroplast and spherical mucocysts. A number of related Euglena species are similar in size, bear one or two stellate plastids and possess spherical or spindle-shaped mucocysts. We conducted morphological and molecular studies on taxa in the subgenus Euglena (all of which bear stellate chloroplasts) and compared this to genera in the subgenus Calliglena (non-stellate chloroplasts). Morphologically the strains in subgenus Euglena were very similar, except for chloroplast number and mucocyst shape. The E. stellata group has one chloroplast and a distinctive spindle-shaped mucocyst; the E. geniculata group has two chloroplasts and spherical mucocysts; the E. viridis group has one chloroplast and spherical mucocysts. Molecular analyses using SSU and LSU rDNA demonstrated that the subgenus Euglena is not monophyletic. The combined SSU/LSU trees provide strong support for a stellate clade (subgenus Euglena ), but one strain of E. viridis diverges at the base of the Euglena/Calliglena lineage. Multiple subclades are found within the main stellate clade. E. tristellata forms a separate divergence and four E. stellata strains form a single, well-supported subclade. Two E. viridis strains are among the E. geniculata group clade, while six others form two separate, but well-supported clades. This study demonstrates that the type species, E. viridis , is paraphyletic and will need to be redefined.