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The Dicranaceae have been classified as one of the largest, most heterogeneous families of the moss subclass Dicranidae. Circumscriptions of the family have varied, with some studies excluding selected subfamilies and recognizing them at the familial rank, whereas others have retained a broader familial concept. As well, classifications have varied in their generic circumscriptions of the subfamilies. Chloroplast DNA sequence data (trnL(UAA)—trnF(GAA) and rps4) were used to examine the monophyly of the family and phylogenetic relationships among the subfamilial and generic taxa. Special emphasis was given to subfamily Dicranoideae, with 18 of the 23 genera sampled. Seventy-four trnL-F and rps4 sequences formed a matrix of 1161 aligned base pairs (bp). Phylogenetic analyses using MP and ML criteria were based on 983 bp (333 parsimony informative) after ambiguous data were removed. Our results support the following inferences: 1) Dicranaceae as traditionally defined are polyphyletic; 2) subfamilies Campylopodioideae, Dicranelloideae, Rhabdoweisioideae, and Trematodontoideae are excluded from a robust monophyletic concept of the Dicranaceae; 3) subfamily Dicranoideae is polyphyletic unless 16 genera are excluded from the subfamily, seven of which are transferred to the Rhabdoweisiaceae; 3) subfamily Paraleucobryoideae is polyphyletic with Brothera resolved in the Leucobryaceae clade and Paraleucobryum nested within subfamily Dicranoideae; 4) Dicnemonaceae (including Mesotus) and Wardiaceae are nested within traditional members of the Dicranaceae; and 5) four clades—Dicranoideae, Mesotoideae, Dicranoloma group plus Wardia, and the Leucoloma group, form a robust monophyletic taxon, considered here as a restricted concept of Dicranaceae (sensu stricto). This circumscription excludes 18 genera that have previously been included in the Dicranaceae.
The tribe Thomsonieae (Araceae) comprise two genera of palaeotropical herbs: Amorphophallus with approximately 200 species and Pseudodracontium with seven species. A phylogenetic study of the Thomsonieae is presented. Nucleotide sequences of the chloroplast gene matK and the trnL intron were analyzed for 46 Amorphophallus species, two Pseudodracontium species and six outgroups using maximum parsimony. The two data sets were analyzed separately and in combination. Amorphophallus can be divided into five major well-supported clades, which provide new hypotheses on infrageneric relationships. The branches connecting these clades, however, are poorly supported. Results are compared with previous morphology-based divisions of Amorphophallus. Remarkable results from this study are the monophyly of the African Amorphophallus species and the nesting of Pseudodracontium within Amorphophallus, making Amorphophallus in the traditional sense paraphyletic to Pseudodracontium.
Previous taxonomic treatments of the family Zosteraceae in Australia/New Zealand have recognized Heterozostera tasmanica (monotypic) and four Zostera species all belonging to subgenus Zosterella: Z. capricorni, Z. muelleri, Z. mucronata, Z. novazelandica. Zostera has always been taxonomically problematic in Australia, where researchers have expressed difficulty with species recognition due to vague or inconsistent morphological characters. There also has been a lack of agreement on generic (notably the distinctness of Heterozostera) and subgeneric delimitation. Recent anatomical, developmental, and molecular studies urge a reevaluation of relationships in the family. To clarify the taxonomy of Zosteraceae, we investigated interspecific phylogenetic relationships focusing on Australian species of subgenus Zosterella. We examined material comprising all genera of Zosteraceae (Heterozostera, Nanozostera, Phyllospadix, Zostera), six/seven species of Zostera subgenus Zosterella (including all Australian/New Zealand species), and one of four species of Zostera subgenus Zostera. We conducted phylogenetic analyses of morphological data and DNA sequences from nuclear (ITS) and plastid (trnK intron, rbcL) genomes. Our results indicate two major clades (highly divergent at both morphological and molecular levels) and two subclades (with low morphological and molecular divergence) within Zosteraceae. Little morphological and molecular variation was observed among representatives within the clade of Australian/New Zealand members of subgenus Zosterella, and none provided cladistic support for taxa recognized formerly as separate species. We recommend that Zosteraceae comprise two genera (Phyllospadix, Zostera) with the latter subdivided into three subgenera (Zostera, Zosterella, Heterozostera). Furthermore, Australian/New Zealand representatives of Zostera subgenus Zosterella should be merged within a single species (Z. capricorni) to reflect the inability of morphological or molecular data to effectively delimit additional species in this group.
A new holomycotrophic species, Thismia taiwanensis (Burmanniaceae), is described and illustrated from Kaohsiung County, south-central Taiwan. It most closely resembles the Japanese species T. abei and T. tuberculata, but is distinct as the flowers are subsessile, the outer surface of the perianth tube is smooth, the outer tepals bear very short solitary appendages, and the inner tepals bear longer solitary appendages. This represents the first record of both Thismia and Burmanniaceae tribe Thismieae in China.
A cladistic analysis using parsimony was carried out, including all six species of Paspalum sect. Pectinata plus an outgroup composed of six species of Paspalum subg. Ceresia, six additional Paspalum species belonging to different taxonomic groups, and two extrageneric taxa. The analysis was based on 65 morphological characters and was performed using both equal weights and implied weights. In all resulting cladograms, a well supported clade corresponding to Paspalum sect. Pectinata appears, with either P. ceresia or a clade comprising P. ceresia, P. stellatum, and P. eucomum as sister group. Within the Pectinata-clade P. lanciflorum emerges consistently as sister taxon to the remainder of the section, whereas the phylogenetic relationship among them is poorly resolved. The inclusion of Paspalum sect. Pectinata within a weakly supported subgenus Ceresia is confirmed. Incidentally, some doubt is thrown about the validity of the currently accepted circumscription of Paspalum subg. Ceresia, since the inclusion of P. humboldtianum and P. polyphyllum within it is not supported by our data.
The phylogenetic positions of the genera Beesia and Eranthis were investigated with respect to seven species in two genera, representing the tribe Actaeeae and five species in five other Ranunculaceae genera as outgroup. Maximum parsimony analyses were performed separately on nuclear ribosomal DNA ITS, plastid trnL-F, and combined DNA sequence data. In these analyses the positions of both Beesia and Eranthis were well supported within the tribe Actaeeae by each analysis and Beesia calthifolia was sister to Anemonopsis macrophylla on a strongly supported clade. Tribe Actaeeae is redefined to include Actaea, Anemonopsis, Beesia, and Eranthis.
A previous analysis of two chloroplast gene regions, rbcL and the rps16 intron, showed a clear phylogenetic pattern in Gunnera. However, these regions were not informative enough to completely resolve the phylogeny. In this study the nuclear ITS region was sequenced for 24 specimens representing 22 species of Gunnera. 223 characters out of 819 were informative and supported the same monophyletic groups as the chloroplast gene regions. Because of its greater information content, the ITS region identified additional well-supported clades. In an analysis based on the three gene regions together, 272 characters out of 3154 were informative. The results show that the South American annual, G. herteri is sister to all other species. The African G. perpensa is well-supported as sister to the remaining species, which form two well-defined clades, one with the Malayan G. macrophylla as sister to subgenus Milligania from New Zealand and Tasmania. In the other clade, the South American subgenus Misandra is sister group to subgenus Panke from South America and Hawaii. Within Panke the two Hawaiian species form a sister group to the American species. The morphological classification of Schindler, together with some biogeographical and morphological features, is discussed.
The trnL (UAA) intron and ITS1 region were sequenced to investigate relationships among the five genera of Vitaceae present in Australia relative to Vitis. Congruent results were obtained between separate and combined data sets, with all major clades being shared among trees. All bootstrap consensus trees obtained from single sequences or combined analysis strongly suggest that Cissus is polyphyletic, corroborating the morphological inconsistencies reported previously. Cissus opaca and Clematicissus angustissima consistently grouped in a common clade. A further four taxa (C. antarctica, C. hypoglauca, C. oblonga, and C. sterculiifolia) also grouped within a clade disjunct from the main Cissus clade. Our results suggest that these five species currently classified as Cissus should be segregated from the genus. Of further interest is the close relationship between Cayratia and Tetrastigma. Overall, the results presented provide new insights into the relationships within a number of Vitaceae genera and suggest directions for future studies.
Myrceugenia fernandeziana (Myrtaceae) is a dominant, apparently anemophilous, tree endemic to Masatierra Island in the Juan Fernandez archipelago. These islands provide a natural laboratory for investigations of relationships within and among populations of threatened/vulnerable species. Leaf morphology was used to compare 25 populations from a diversity of habitats. Allozyme analyses were conducted for a subset of 15 of these populations. Leaf shape, described by elliptic Fourier analyses, allows recognition of distinct subsets of populations (also recognized by allozymes) and a number of leaf characters show evidence of geographically-based (southeast to northwest) clinal variation on the island. Morphological distances among populations are not correlated with geographic distances but allozyme distances among populations are correlated with geographic distances. There is a positive, but not statistically significant, correlation between allozyme and morphological distances. In addition, there is no apparent connection between patterns of allozyme or morphological differences and the habitats in which the populations occur. These results are not consistent with models based on localized adaptation, suggest panmixis, perhaps based on anemophily and broad dispersal, and suggest that conservation strategies must be based on sampling multiple populations within and among habitat types.
To understand flower morphological evolution in Gentianaceae-Swertiinae, we studied generic relationships using trnL intron, matK, and nuclear ITS sequences of a total of 13 genera and 59 species of the subtribe. The phylogenetic incongruencies between the chloroplast and nuclear genes are likely to be the result of long branch attraction. The East Asian Megacodon and Latouchea and the eastern North American Bartonia and Obolaria were determined as the most basal genera, and several well-supported subgroups were revealed. Swertia, Lomatogonium, and Gentianella s. l. were highly polyphyletic and the position of Veratrilla and several species was ambiguous. The main flower types found in Swertiinae can be transformed into each other by simple developmental variation in proportion. This apparently happened several times during the evolution of Swertiinae and, in conjunction with other homoplastic characters, explains the difficulty of recognizing generic limits and the mosaic pattern of character distribution. Phylogenetic relationships, extant distribution ranges, and a preliminary molecular clock approach led to the hypothesis that the last common ancestor of the Swertiinae lived approximately 15 mya, and that an exchange of lineages between East Asia and North America happened frequently from the time of origin until only recently.
Gasteranthus (Gesneriaceae; Beslerieae) includes 38 species restricted to the New World tropics. The genus has a limited distribution in Central and South America, with a center of diversity in western Ecuador. This study investigates cladistic relationships among species of Gasteranthus using floral and vegetative morphological data. Corolla shape in Gasteranthus ranges from funnelform to urceolate and ventricose, with additional variation in corolla spur and corolla ventral pouch morphology. Our analysis revealed that there were likely single primary innovations of the three major corolla types, although only one of the three types forms a monophyletic group. Apparently, ventricose corollas were derived from funnelform corollas and urceolate corollas were derived from ventricose corollas, making the funnelform and ventricose corolla groups both paraphyletic. Spurs have been lost twice in Gasteranthus. Conspicuousness of stomatal clusters is quite homoplastic given the phylogenetic hypothesis.
Sequences of nuclear ribosomal DNA ITS and ETS regions were used to examine phylogenetic relationships of Syringa and Ligustrum. Twenty-seven samples were included in parsimony analyses, representing all major groups of these two genera. Two species of Fraxinus and one species of Jasminum were also included in analyses for rooting purposes. In the resulting phylogenetic hypothesis series Syringa (Syringa) diverges first and is followed by the monotypic series Pinnatifoliae (Syringa), which is sister to a clade containing the remaining species of Syringa and species of Ligustrum. However, this sister relationship is weakly supported. Our results support the recognition of monophyletic groups corresponding to subgenus Ligustrina, and series Syringa, Pubescentes, and Villosae, and suggest that Ligustrum as a monophyletic group is derived from within Syringa, such that Syringa as traditionally circumscribed is paraphyletic. Forcing Syringa to be monophyletic entails nine extra steps, which is significant, as judged by the Templeton test. Parasyringa sempervirens is phylogenetically embedded within the Ligustrum clade, supporting its placement in Ligustrum. Berries are a synapomorphy of Ligustrum species, and the dehiscent berry of Parasyringa sempervirens is likely to be an evolutionary reversal towards a capsule, which is characteristic of Syringa. The evolution of berries might have contributed to an accelerated rate of speciation in Ligustrum.
Mutisieae are considered one of the most important tribes for understanding the systematics and evolution of the Asteraceae because of their basal position within the family. The tribe is extremely diverse morphologically and biogeographically, consisting of 84 genera and 900 species. Despite extensive studies using macromorphological, palynological, and molecular approaches, tribal and subtribal delimitation and relationships of Mutisieae remain controversial. Phylogenetic analyses of DNA sequences of the chloroplast gene ndhF were performed using 53 species representing 31 genera of Mutisieae, 11 genera from the remaining five tribes of Cichorioideae, and five genera from the Asteroideae. The ndhF phylogeny of the Mutisieae provides insights into the circumscription and relationships of the tribe, subtribes, and taxonomic placement of several morphologically anomalous genera. The ndhF tree indicates that Mutisieae are polyphyletic. The subtribes Gochnatiinae and Mutisiinae are not monophyletic, whereas the Nassauviinae are monophyletic. The majority of the South American genera and the Chinese genus Nouelia of the Gochnatiinae are positioned in a basal position in the Mutisieae. Furthermore, intergeneric relationships are resolved in several cases.
A morphometric analysis was conducted of Echinacea Moench (Asteraceae) to measure variation between native populations for taxonomic purposes. Data were collected from living and herbarium specimens. From a matrix of 321 specimens by 74 characters, a pair-wise distance matrix was computed using Gower's coefficient. Cluster strategies were explored from the distance matrix. MODECLUS clustering separated the data into two clusters, and a flexible agglomerative clustering method separated the data into the same two clusters, which were broken into four sub-clusters. Canonical discriminant analysis gave significant support for the two- and the four-cluster solutions. Canonical discriminant analysis also showed support for eight smaller clusters identified using McGregor's 1968 classification. We recognize two subgenera and four species: Echinacea subg. Echinacea contains only E. purpurea; Echinacea subg. Pallida contains E. atrorubens, E. laevigata, and E. pallida. The revised varieties are as follows: E. atrorubens var. atrorubens, E. atrorubens var. neglecta, E. atrorubens var. paradoxa, E. pallida var. angustifolia, E. pallida var. pallida, E. pallida var. sanguinea, E. pallida var. simulata, and E. pallida var. tennesseensis. A cladistic analysis was done on the four species. In the most parsimonious solution, E. purpurea was basally divergent to a clade of the other three species (70% bootstrap value), and all four were distinguishable by at least one apomorphy. A key to Echinacea taxa is provided, which should be valuable given the pharmaceutical and horticultural importance of Echinacea.
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