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Monocot systematics: a combined analysis
... La estructura de las ceras epicuticulares caracteriza a grandes grupos de monocotiledóneas. En éstos se ha observado en abundancia dos tipos de ceras, las que tienen forma de varillas agregadas en forma longitudinal y aquellas orientadas en forma de placas paralelas, estas últimas encontradas en la familia Nolinaceae (Chase et al., 1995). ...
... waxes characterizes large groups of monocots. In these has been observed in abundance two types of waxes, which have added rodlike longitudinally oriented and those in the form of parallel plates, the latter found in the family Nolinaceae (Chase et al., 1995). ...
En el presente trabajo se analizó la morfo-anatomía foliar de sotol Dasylirion cedrosanum Trel., especie perenne del Norte de México y Sur de Estados Unidos de América. Esta especie es de gran importancia económica y social en las zonas áridas y semiáridas de México. El objetivo fue observar las diferencias en el desarrollo morfo-anatómico de la hoja a diferentes edades. Para el estudio morfológico se tomaron muestras de plantas de 6, 30, 60 y 84 meses de edad. Se colectaron porciones medias del eje longitudinal de las hojas en las edades previamente establecidas y se fijaron en FAA. Las muestras se procesaron con los métodos clásicos para la obtención de láminas permanentes. La epidermis foliar está formada por células unistratas de forma isodiamétrica, los estomas son de tipo paracítico, la densidad estomática, el índice estomático muestra valores entre 14.0–23.3% y la densidad de células estomáticas oscila entre 46.8-65, estomas mm-2, las superficies adaxial y abaxial no exhiben dominancia una sobre otra. El parénquima en empalizada se incrementa hasta 62% con la edad; el mesofilo es isolateral con braquiesclereidas que se prolongan desde las bandas vasculares. Los haces vasculares son bandas de tres órdenes sobre la que existen cristales prismáticos.
... La estructura de las ceras epicuticulares caracteriza a grandes grupos de monocotiledóneas. En éstos se ha observado en abundancia dos tipos de ceras, las que tienen forma de varillas agregadas en forma longitudinal y aquellas orientadas en forma de placas paralelas, estas últimas encontradas en la familia Nolinaceae (Chase et al., 1995). ...
... waxes characterizes large groups of monocots. In these has been observed in abundance two types of waxes, which have added rodlike longitudinally oriented and those in the form of parallel plates, the latter found in the family Nolinaceae (Chase et al., 1995). ...
Abstract: In this paper was analyzed the foliar morpho-anatomy of Sotol Dasylirion cedrosanum Trel., perennial species northern Mexico and southern United States. This species is of great economic and social importance in arid and semiarid areas of Mexico. The objective was to observe the differences in morphological and anatomical leaf development at different ages. For morphological study plant samples 6, 30, 60 and 84 months of age they were taken. The middle portions of the longitudinal axis of leaves were collected at ages previously established and fixed in FAA. The samples were processed with conventional methods to obtain permanent slides. The leaf epidermis consists unistrate cells isodiametric form, the stomata are paracytic type, stomatal density, stomatal index shows values between 14.0-23.3% and stomatal cells density ranging from 46.8-65, stomata mm-2 , the adaxial and abaxial surfaces do not exhibit dominance over one another. The palisade parenchyma up to 62% increases with age; the mesophyll is isolateral with brachisclereids extending from vascular bands. The vascular bundles are bands of three orders on which there prismatic crystals.
... Taxon sampling followed the multilocus data sets of Chase et al. (1995 Chase et al. ( , 2000 Chase et al. ( , 2006). These data included 124 species representing all 11 orders of monocots and Dasypogonaceae (Givnish et al., 2006); all extant monocot families circumscribed by APG III (2009) were represented except for three families in Alismatales (Ruppiaceae, Posidoniaceae and Scheuchzeriaceae). ...
... In most cases the DNA used for amplification was the same as was used in previous molecular phylogenetic studies of the monocots (File S1; Chase et al., 1995 Chase et al., , 2000 Chase et al., , 2006). Other taxa represented the same genus or family when DNA accessions were unavailable and/or did not amplify; estimations of familial relationships using similar procedures have shown that such substitutions have not had adverse effects on phylogenetic studies at higher taxonomic levels as these families are monophyletic (Qiu et al., 1999; Soltis et al., 2000). ...
Resolution of evolutionary relationships among some monocot orders remains problematic despite the application of various taxon and molecular locus sampling strategies. In this study we sequenced and analysed a fragment of the low-copy, nuclear phytochrome C (PHYC) gene and combined these data with a previous multigene data set (four plastid, one mitochondrial, two nuclear ribosomal loci) to determine if adding this marker improved resolution and support of relationships among major lineages of monocots. Our results indicate the addition of PHYC to the multigene dataset increases support along the backbone of the monocot tree, although relationships among orders of commelinids remain elusive. We also estimated divergence times in monocots by applying newly evaluated fossil calibrations to our resolved phylogenetic tree. Inclusion of early-diverging angiosperm lineages confirmed the origin of extant monocots c. 131 Mya and strengthened the hypothesis of recent divergence times for some lineages, although current divergence time estimation methods may inadequately model rate heterogeneity in monocots. We note significant shifts in diversification in at least two monocot orders, Poales and Asparagales. We describe patterns of diversification in the context of radiation of other relevant plant and animal lineages. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, ●●, ●●–●●.
... However, it is also possible that the additional taxa in Chase et al. (2000) caused a greater level of resolution. The major issue for this analysis is that our results do not contradict those of Chase et al. (2000), and those of the latter are highly similar to previous analyses of rbcL alone for the monocots (Duvall et al., 1993; Chase et al., 1995). The tree illustrated here (Fig. 3B) is in agreement with the strict consensus tree of Chase et al. (2000), and the lack of resolution is again centred around the shortest branches (all of these have 10–16 steps whereas those branches nearby with jackknife support all have 21 or more steps; Fig. 3B), leading us to conclude that obtaining a robust assessment of the branching order simply requires more data. ...
... These two families are, in turn, sister to a clade (73%) that includes Iridaceae, Xeronemataceae, Anthericaceae, Behniaceae, Agavaceae, Laxmanniaceae, Asphodeliaceae , Xanthorrhoeaceae, Amaryllidaceae, Alliaceae, Asparagaceae, Hyacinthaceae and Convallariaceae. After the analysis was completed, it was noticed that the taxon labelled in the matrix as 'Luzuriaga' fell within Asparagales; the rbcL sequence alone for this terminal appears in Liliales (as in Chase et al., 1995). The identity of the specimen used for the atpB and 18S rDNA sequences was originally determined to be Luzuriaga latifolia (W. ...
A phylogenetic analysis of a combined data set for 560 angiosperms and seven outgroups based on three genes, 18S rDNA (1855 bp), rbcL (1428 bp), and atpB (1450 bp) representing a total of 4733 bp is presented. Parsimony analysis was expedited by use of a new computer program, the RATCHET. Parsimony jackknifing was performed to assess the support of clades. The combination of three data sets for numerous species has resulted in the most highly resolved and strongly supported topology yet obtained for angiosperms. In contrast to previous analyses based on single genes, much of the spine of the tree and most of the larger clades receive jackknife support ≥50%. Some of the noneudicots form a grade followed by a strongly supported eudicot clade. The early-branching angiosperms are Amborellaceae, Nymphaeaceae, and a clade of Austrobaileyaceae, Illiciaceae, and SchiÍsandraceae. The remaining noneudicots, except Ceratophyllaceae, form a weakly supported core eumagnoliid clade comprising six well-supported subclades: Chloranthaceae, monocots, Winteraceae/Canellaceae, Piperales, Laurales, and Magnoliales. Ceratophyllaceae are sister to the eudicots. Within the well-supported eudicot clade, the early-diverging eudicots (e.g. Proteales, Ranunculales, Trochodendraceae, Sabiaceae) form a grade, followed by the core eudicots, the monophyly of which is also strongly supported. The core eudicots comprise six well-supported subclades: (1) Berberidopsidaceae/Aextoxicaceae; (2) Myrothamnaceae/Gunneraceae; (3) Saxifragales, which are the sister to Vitaceae (including Leea) plus a strongly supported eurosid clade; (4) Santalales; (5) Caryophyllales, to which Dilleniaceae are sister; and (6) an asterid clade. The relationships among these six subclades of core eudicots do not receive strong support. This large data set has also helped place a number of enigmatic angiosperm families, including Podostemaceae, Aphloiaceae, and Ixerbaceae. This analysis further illustrates the tractability of large data sets and supports a recent, phylogenetically based, ordinal-level reclassification of the angiosperms based largely, but not exclusively, on molecular (DNA sequence) data.
... Setting aside this probable artefact here, members of Thismiaceae are therefore inferred to be monophyletic at the current taxon sampling, and are phylogenetically distinct from Burmanniaceae. Our results are consistent with recognizing Burmanniaceae and Thismiaceae as two distinct lineages in the order, conflicting withJonker (1938), Maas et al. (1986), APG (2003), Caddick et al. (2002a, but in agreement withChase et al. (1995),Merckx et al. (2006), Woodward (2007,Merckx and Smets (2014),Cheek et al. (2018),Lam et al. (2016;2018),Shepeleva et al. (2020) andLin et al. (2022). ...
Premise:
Species in Thismiaceae can no longer photosynthesize, and instead obtain carbon from soil fungi. Here we infer Thismiaceae phylogeny using plastid genome data, and also characterize the molecular evolution of this genome.
Methods:
We assembled five Thismiaceae plastid genomes from genome skimming data, adding to previously published data for phylogenomic inference. We investigated plastid-genome structural changes, considering locally colinear blocks (LCBs). We also characterized possible shifts in selection pressure in retained genes by considering changes in ω, the ratio of non-synonymous to synonymous changes.
Key results:
Thismiaceae experienced two major pulses of gene loss around the early diversification of the family, with subsequent scattered gene losses across descendent lineages. In addition to massive size reduction, Thismiaceae plastid genomes experienced occasional inversions, and there likely were two independent losses of the plastid inverted repeat (IR) region. Retained plastid genes remain under generally strong purifying selection (ω << 1), with significant and sporadic weakening or strengthening in several instances. The bifunctional trnE-UUC gene of Thismia huangii likely retains a secondary role in heme biosynthesis, despite a probable loss of functionality in protein translation. Several cis-spliced group IIA introns are retained, despite the loss of the plastid intron maturase, matK.
Conclusions:
We infer that most gene losses in Thismiaceae occurred early and rapidly, following the initial loss of photosynthesis in its stem lineage. As a species-rich, fully mycoheterotrophic lineage, Thismiaceae provides a model system for uncovering the unique and divergent ways in which plastid genomes evolve in heterotrophic plants. This article is protected by copyright. All rights reserved.
... In addition, they had paired interpetiolar tendrils, mostly spinous stems, unisexual flowers with six tepals, and either six fertile stamens or staminodes in pistillate flowers, umbellate inflorescence, including fleshly berries. This family has a long nomenclature history and has been placed in various positions, such as in Liliales sensu lato (s.l.) (Hutchinson, 1979;Thorne, 1983;Goldberg, 1989;Cronquist, 1991), Dioscoreales (Dahlgren & Clifford, 1982;Thorne, 1992) or Asparagales (Hurber, 1969 (Chase et al., 1995;Patterson & Givnish, 2002;Fay et al., 2006;Kim et al., 2013;Petersen et al., 2013). ...
Smilacaceae is known as a taxa with wide phenotypic variation and their taxonomical complexities remain unsolved. The three species of Smilacaceae housed in Java, are given nomenclature history. Since all the potential sources of original material have been investigated, but nothing has been identified, three neotypifications were designated here. Herbarium specimen of Koorders 34990β in Herbarium Bogoriense (BO) was chosen as the neotype of Smilax klotzschii. The de Groot & Wehlburg RD52 herbarium specimen in BO was chosen as the neotype of S. nageliana and Blume 463 herbarium specimen in L was chosen as the neotype of S. odoratissima.
... Certain similarities can be observed in other highly diverse classes of organisms, such as Magnoliophyta (flowering plants, 300,000 species 52 ). Loss of the plant ancestral telomere DNA sequence, (TTT AGG G) n , has been reported in numerous flowering plants, in which the sequences were replaced by alternative motifs 10,53-57 , and high diversity in telomere repeats was observed in two species-rich orders, Asparagales, the largest order within the monocotyledons, consisting of around 30,000 species [58][59][60][61] , and Lamiales, consisting of 23,000 species 55 . Besides, numerous plant orders possess unknown telomere sequences 10 . ...
Telomeres are protective structures at the ends of eukaryotic chromosomes, and disruption of their nucleoprotein composition usually results in genome instability and cell death. Telomeric DNA sequences have generally been found to be exceptionally conserved in evolution, and the most common pattern of telomeric sequences across eukaryotes is (TxAyGz)n maintained by telomerase. However, telomerase-added DNA repeats in some insect taxa frequently vary, show unusual features, and can even be absent. It has been speculated about factors that might allow frequent changes in telomere composition in Insecta. Coleoptera (beetles) is the largest of all insect orders and based on previously available data, it seemed that the telomeric sequence of beetles varies to a great extent. We performed an extensive mapping of the (TTAGG)n sequence, the ancestral telomeric sequence in Insects, across the main branches of Coleoptera. Our study indicates that the (TTAGG)n sequence has been repeatedly or completely lost in more than half of the tested beetle superfamilies. Although the exact telomeric motif in most of the (TTAGG)n-negative beetles is unknown, we found that the (TTAGG)n sequence has been replaced by two alternative telomeric motifs, the (TCAGG)n and (TTAGGG)n, in at least three superfamilies of Coleoptera. The diversity of the telomeric motifs was positively related to the species richness of taxa, regardless of the age of the taxa. The presence/absence of the (TTAGG)n sequence highly varied within the Curculionoidea, Chrysomeloidea, and Staphylinoidea, which are the three most diverse superfamilies within Metazoa. Our data supports the hypothesis that telomere dysfunctions can initiate rapid genomic changes that lead to reproductive isolation and speciation.
... The Eriocaulaceae are a monophyletic family classified in Poales (Chase & al., 1995;Linder & Kellogg, 1995;Stevenson & Loconte, 1995;Givnish & al., 1999;Giulietti & al., 2000;Bremer, 2002;APG III, 2009). They are easily characterized by graminoid leaves, small unisexual flowers grouped in dense capitula, a 3-or 2-locular ovary (with a single pendulous ovule per locule), and having spiraperturate p ollen grains (Giulietti & al., 1995(Giulietti & al., , 2000). ...
The Eriocaulaceae are easily recognized because of the small unisexual flowers in long–pedunculate heads and spiraperturate pollen grains. Their monophyly has never been disputed but internal relationships within the family have not been broadly explored and genera are typically distinguished by few floral characters. Here, we present the first comprehensive phylogenetic study of Eriocaulaceae based on individual and combined molecular datasets, including the plastid psbA–trnH and trnL–F and the nuclear ITS. Results are largely congruent among DNA regions and support the internal dichotomy between the two subfamilies: Eriocauloideae and Paepalanthoideae. Eriocaulon and Leiothrix are strongly supported as monophyletic, whereas Paepalanthus, Blastocaulon, and Syngonanthus are not monophyletic. The phylogenetic nature of Actinocephalus is not resolved and Lachnocaulon, Mesanthemum, Philodice, Rondonanthus, and Tonina (monospecific) are represented in our analyses by a single species each. Based on our results, we suggest two principal generic realignments in the family. The first is the division of Syngonanthus in two genera: Syngonanthus s.str., including Philodice (conserving the former name against the latter), and Comanthera, which is being reinstated to include two sections segregated from Syngonanthus: S. sect. Eulepis and S. sect. Thysanocephalus. The second realignment suggested is the amalgamation of Actinocephalus, Blastocaulon, Lachnocaulon, and Tonina into Paepalanthus s.l. Three 'stat. nov.' within Paepalanthus are published (P. ser. Leptocephali, P. ser. Rosulati, and P. ser. Dimeri).
... Juntamente con Arecales, Commelinales y Poales (APG II 2003, APG III 2009, las Zingiberales, integran el clado de las monocotiledóneas comelinoides, tratado como monofilético sobre la base de la morfología y la biología molecular, como las secuencias rbcL, rbcL, atpB y 18S (Dalgren & Rasmussen 1983, Kress 1990, Chase et al. 1995, Duvall 1993, Kress 1995, Freire Fierro 2004. Judd et al. (2002) las señalan como el orden más avanzado de las Monocotiledóneas y consideran ocho familias: Cannaceae, monotípica, con unas 20-22 especies, predominantemente neotropicales (Ciciarelli & Rolleri 2008); Costaceae, con unos 7 géneros y cerca de 100 especies de zonas tropicales de Asia, África y América; Heliconiaceae, también monotípica, con unas 7 especies neotropicales; Lowiaceae, con unas 15 especies agrupadas sub Orchidanta N. E. Brown, que crecen desde el S de China hasta Borneo; Musaceae, con 2 géneros y 35 especies; Marantaceae, con 31 géneros y unas 550 especies, predominantemente neotropicales; Strelitziaceae, con 3 géneros y unas 7-8 especies de las regiones tropicales y sub-tropicales de América y África, y Zingiberaceae, la familia más numerosa, con unos 50 géneros y unas 1400 especies originarias del paleotrópico (Indonesia y Malasia), introducidas o naturalizadas en Norteamérica, México, Antillas, Mesoamérica, Sudamérica, Asia, África y Australia. ...
Se presenta un estudio, efectuado con microscopía de luz y electrónica de barrido, de la morfología del polen de representantes nativos, naturalizados y cultivados de Zingiberales, que crecen en humedales rioplatenses y selvas subtropicales del NE de la Argentina. Las especies analizadas son Canna glauca y C. índica (Cannaceae), Heliconia psittacorum, H. brasiliensis y H. rostrata (Heliconiaceae), Thalia geniculata y Maranta leuconeura (Marantaceae), Musa acuminata × balbisiana y M. × paradisiaca (Musaceae), Strelitzia nicolai y S. reginae (Strelitziaceae), Alpinia zerumbet, Hedychium coronarium y H. gardnerianum (Zingiberaceae). Se encontraron granos esferoidales, oblados, apolares y heteropolares, inaberturados, con exinas delgadas e intinas gruesas con dos capas. Los tipos de ornamentación hallados son: equinados, típicos o micro-equinados, de Cannaceae, Heliconiaceae y algunas Zingiberaceae, muriformes, típicos o micro-muriformes (rugados, rugulados, areolados) y lisos, presentes en Musaceae, Strelitziaceae y Zingiberaceae. El polen de Thalia geniculata, los granos rugados de Musaceae y los areolados de Strelitziaceae se describen por primera vez, y se analizan rasgos, como las hebras de elastina y el polenkit, que aportan caracteres actualizados para el conocimiento de los taxones seleccionados, poco difundidos desde el punto de vista palinológico.
... updated), are considered to be a natural group. The monophyly of Bromeliaceae has been consistently confirmed in several studies (Stevenson & Loconte, 1995;Chase et al., 1995Chase et al., , 2000Givnish et al., 2011). Givnish et al. (2007) suggested that a recent diversification of modern lineages of Bromeliaceae occurred c. 19 Mya. ...
We expand the genome size (GS) database for Bromeliaceae, specifically for subfamily Pitcairnioideae, and verify whether GS can provide information on the diversification of the five genera in this subfamily. We also provide a phylogenetic perspective on GS evolution in the subfamily and reconstruct the ancestral state for this character. We show that the evolutionary path of GS from the origin of angiosperms to the origin of Pitcairnioideae agrees with the proportional model of GS evolution. Furthermore, we propose that the high phenotypic diversity that is found across Bromeliaceae and that is well represented in Pitcairnioideae is both correlated with high rates of GS evolution of the species and associated with a short period of diversification. The paper also highlights the value of flow cytometry as a rapid and reliable technique for generating GS data which can be analysed in conjunction with other molecular and morphological data to help elucidate patterns of evolution and phylogenetic relationships within this family.
... In addition to broad analyses of major clades of green life, other studies focused on major subclades, including the monocots (e.g. Chase et al., 2006Chase et al., , 2000Chase, Stevenson, Wilkin, & Rudall, 1995;Duvall et al., 1993;Givnish et al., 2006). There were also numerous foundational studies at finer scales-within orders and families of flowering plants (e.g. ...
From restriction site analyses to whole plastid genome sequences, our understanding of green plant (Viridiplantae; ∼. 500,000 extant species) evolutionary relationships over the past three decades has largely been informed by analyses of the plastid genome. The plastid genome has informed studies ranging from population genetics to phylogenetics, the latter ranging from the intraspecific level to studies of all green plants. Diverse portions of the genome ranging from plastid spacers to entire genomes provide valuable data for plant evolutionary biologists. Recent phylogenetic analyses using whole plastid genomes sampled from over 2000 species representing all major groups of green plants have both solidified our understanding of relationships and highlighted the few key nodes in plant evolutionary history that remain unresolved. Likewise, detailed large-scale analyses of plastomes across angiosperms reinforce firmly supported nodes but fail to resolve a handful of remaining questionable relationships. The long history of plastid phylogenetics will serve as a reference point as scientists continue to expand beyond the plastid genome and include more nuclear and mitochondrial data in their analyses. These comparisons are crucial in that recent studies indicate some discordance between nuclear and plastid gene trees both across green plants as a whole and within angiosperms. Rather than being a source of concern, these discordances point to the complex and intriguing one-billion-year evolutionary history of the green plant clade, a clade that is foundational to life on Earth.
... are a widely recognised family of mycoheterotrophic flowering plants (Chase et al. 1995, Chase et al. 2000, Woodward et al. 2007, Merckx 2013, Merckx & Smets 2014, Truòng et al. 2014. DNA analyses have indicated that the five genera typically included in tribe Thismieae, namely, Thismia Griffith (1845: 221), Afrothismia Schlechter (1906: 138), Oxygyne Schlechter (1906: 140), Haplothismia Airy Shaw (1952: 277) and Tiputinia Berry & Woodward (2007: 158), all fall outside Burmanniaceae (Merckx & Smets 2014), but their specific placement within the order Dioscoreales remains unclear (Merckx et al. 2009, Merckx & Smets 2014. ...
Thismia nigricoronata is described as a new species in family Burmanniaceae. Both morphological and phylogenetic analyses indicate that this new Lao endemic is allied to T. taiwanensis in section Glaziocharis, and it can be differentiated on the basis of its longer vestigial stem leaves, reflexed free outer perianth lobes and ornamented, vibrantly coloured outer surface of the perianth tube. The infrageneric taxonomy of Thismia is reviewed, the genera Geomitra and Scaphiophora are officially reduced to sectional status in Thismia, and all species are enumerated in systematic order. A key to all currently accepted subgenera, sections and subsections is presented to facilitate further examination of their phylogenetic integrity in light of apparent conflict between the traditional morphology-based system and the emerging DNA-based classification.
... Scale bars are 5 µm in a, 1 µm in b and c, 500 nm in d and g, 100 nm in e, 2 µm in f (Hamann 1966) and in the periplasmodium in Haemodoraceae (e.g., Lachnanthes;Simpson 1988). Phylogenetic analyses indicate that they are strongly related to the family Commelinaceae (Chase et al. 1995), which suggests that tapetal raphides should be regarded as a taxonomic trait. Formation of tapetal raphides may result from accumulation of excessive amounts of calcium and oxalic acid salts in the anther locules. ...
Calcium oxalate (CaOx) crystals in higher plants occur in five forms: raphides, styloids, prisms, druses, and crystal sand. CaOx crystals are formed in almost all tissues in intravacuolar crystal chambers. However, the mechanism of crystallization and the role of CaOx crystals have not been clearly explained. The aim of this study was to explore the occurrence and location of CaOx crystals in organs of Tinantia anomala (Torr.) C.B. Clarke (Commelinaceae) with special attention to ultrastructural changes in the quantity of tapetal raphides during microsporogenesis. We observed various parts of the plant, that is, stems, leaves, sepals, petals, anthers, staminal trichomes and stigmatic papillae and identified CaOx crystals in all parts except staminal trichomes and stigmatic papillae in Tinantia anomala. Three morphological forms: styloids, raphides and prisms were found in different amounts in different parts of the plant. Furthermore, in this species, we identified tapetal raphides in anthers. The number of tapetal raphides changed during microsporogenesis. At the beginning of meiosis, the biosynthesis of raphides proceeded intensively in the provacuoles. These organelles were formed from the endoplasmic reticulum system. In the tetrad stage, we observed vacuoles with needle-shaped raphides (type I) always localised in the centre of the organelle. When the amoeboid tapetum was degenerating, vacuoles also began to fade. We observed a small number of raphides in the stage of mature pollen grains.
... Ellos sostienen que Dioscoreaceae está muy relacionada con Smilacaceae y ésta última con Petermanniaceae y Liliaceae. Tanto estudios morfológicos (Conran 1989) como de secuencias de ADN (Chase et al. 1995a, 1995b, 2000, Judd et al. 1999, 2002, Rudall et al. 2000, Patterson & Givnish 2002 favorecen la ubicación de Smilacaceae en el orden Liliales. ...
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Se realizó una revisión morfológica de las especies de Smilax de Costa Rica. Tradicionalmente se han aceptado hasta 14 especies. En este trabajo se reconocen 7: Smilax domingensis, S. mollis, S. panamensis, S. spinosa, S. spissa, S. subpubescens y S. vanilliodora. Los siguientes nombres se tratan como sinónimos: Smilax engleriana y S. kunthii de S. domingensis ; S. hirsutior, S. angustiflora y S. candelariae de S. mollis , y S. chiriquensis y S. regelii var. albida de S. vanilliodora . Smilax regelii se excluye como taxon válido y se designa un lectotipo de S. gymnopoda . Se elaboraron claves dicotómicas con características vegetativas y reproductivas (flores y frutos), con base en observaciones de campo y especímenes de herbario. En todas las especies se incluyeron diversos caracteres importantes para la identificación, como el rizoma, el tallo, el tamaño de los tépalos y la variación del color de las bayas a lo largo del desarrollo.
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... Numerous phylogenetic analyses of the monocots conducted over the past 20 years, employing a variety of molecular and morphological characters, have contributed to the development of an increasingly stable taxonomic system for the c. 80-100 conventionally recognized families of monocots (e.g. Duvall et al., 1993;Chase et al., 1995Chase et al., , 2000Stevenson and Loconte, 1995;Neyland and Hennigan, 2003;Davis et al., 2004Davis et al., , 2006Janssen and Bremer, 2004;Tamura et al., 2004;Givnish et al., 2006;Graham et al., 2006;Xiao-Xian and Zhe-Kun, 2007). Current understandings of relationships among major groups are summarized in the APG III system (APG III, 2009), in which 12 major clades of monocots are recognized, 11 of them at the ordinal level, and the remaining group (the family Dasypogonaceae) unassigned to order, but included in the informally recognized commelinid alliance with Commelinales, Zingiberales, Arecales and Poales. ...
Tracing the evolution of one of the most ancient major branches of flowering plants, this is a wide-ranging survey of state-of-the-art research on the early clades of the monocot phylogenetic tree. It explores a series of broad but linked themes, providing for the first time a detailed and coherent view of the taxa of the early monocot lineages, how they diversified and their importance in monocots as a whole. Featuring contributions from leaders in the field, the chapters trace the evolution of the monocots from largely aquatic ancestors. Topics covered include the rapidly advancing field of monocot fossils, aquatic adaptations in pollen and anther structure and pollination strategies and floral developmental morphology. The book also presents a new plastid sequence analysis of early monocots and a review of monocot phylogeny as a whole, placing in an evolutionary context a plant group of major ecological, economic and horticultural importance.
... However, molecular data have yielded varying results regarding the phylogenetic position of Campynemataceae when species of Corsiaceae were included in the analyses. Plastid rbcL sequence data have supported both a sister relationship of Campynemataceae to Liliales (Chase et al. 1995;Janssen and Bremer 2004) as well as a sister relationship to the Melanthiaceae-Smilacaceae-Liliaceae clade with strong bootstrap support (Vinnersten and Bremer 2001). Upon addition of nuclear (18S, 26S), mitochondrial (atp1), and plastid data (atpB, matK, ndhF, and rbcL), Campynemataceae and Melanthiaceae comprise a clade that was sister to the remaining Liliales, but this position showed weak support . ...
The phylogenetic positions of the families Campynemataceae and Corsiaceae within the order Liliales remains unclear. To date, molecular data from the plastid genome of Corsiaceae has been obtained exclusively from Arachnitis, for which alignment and phylogenetic inference has proved difficult. The extent of gene conservation among mycoheterotrophic species within Corsiaceae remains unknown. To clarify the phylogenetic position of Campynemataceae and Corsiaceae within Liliales, functional plastid-coding genes of species representing both families have been analyzed. Examination of two phylogenetic data sets of plastid genes employing parsimony, maximum-likelihood, and Bayesian inference methods strongly supported both families forming a basal clade to the remaining taxa of Liliales. The first data set consists of five functional plastid-encoded genes (matK, rps7, rps2, rps19, and rpl2) sequenced from Corsia dispar (Corsiaceae). The data set included 31 species representing all families within Liliales, as well as selected orders that are related closely to Liliales (10 outgroup species from Asparagales, Dioscoreales, and Pandanales). The second phylogenetic analysis was based on 75 plastid genes. This data set included 18 species from Liliales, representing major clades within the order, and 10 outgroup species from Asparagales, Dioscoreales, and Pandanales. In this latter data set, Campynemataceae was represented by 60 plastid-encoded genes sequenced from herbarium material of Campynema lineare. A large proportion of the plastid genome of C. dispar was also sequenced and compared to the plastid genomes of photosynthetic plants within Liliales and mycoheterotrophic plants within Asparagales to explore plastid genome reduction. The plastid genome of C. dispar is in the advanced stages of reduction, which signifies its high dependency on mycorrhizal fungi and is suggestive of a loss in photosynthetic ability. Functional plastid genes found in C. dispar may be applicable to other species in Corsiaceae, which will provide a basis for in-depth molecular analyses of interspecies relationships within the family, once molecular data from other members become available.
... The relationship (Tofieldiaceae, (Araceae, core alismatids [+/À Acorus])) has been suggested-but mostly not well supported-by previous analyses (e.g. Chase et al. 1995Chase et al. , 2000Duvall et al., 2006;Graham et al., 2006;Iles et al., 2013), but most phylogenetic analyses yield an alternative resolution: (Araceae, (Tofieldiaceae, core alismatids [+/À Acorus])) (e.g. Qiu et al., 2000Qiu et al., , 2006Qiu et al., , 2010Davis et al., 2004;Tamura et al., 2004;Chase et al., 2006;Givnish et al., 2006;Petersen et al., 2006a;Soltis et al., 2007;Azuma and Tobe, 2011). ...
A phylogenetic analysis of the early branching lineages of the monocotyledons is performed using data from two plastid genes (rbcL and matK), five mitochondrial genes (atp1, ccmB, cob, mttB and nad5) and morphology. The complete matrix includes 93 terminals representing Acorus, the 14 families currently recognized within Alismatales, and numerous lineages of monocotyledons and other angiosperms. Total evidence analysis results in an almost completely resolved strict consensus tree, but all data partitions, genomic as well as morphological, are incongruent. The effects of RNA editing and potentially processed paralogous sequences are explored and discussed. Despite a decrease in incongruence length differences after exclusion of edited sites, the major data partitions remain significantly incongruent. The 14 families of Alismatales are all found to be monophyletic, but Acorus is found to be included in Alismatales rather than being the sister group to all other monocotyledons. The placement is strongly supported by the mitochondrial data, atp1 in particular, but it cannot be explained as an artifact caused by patterns of editing or by sampling of processed paralogues.
... However, use of molecular data, has shed new light on the position of Burmanniaceae among the monocots. In a phylogenetic analysis of 172 monocot rbcL sequences, a Burmannia species was sister to Dioscorea and Tacca (Chase et al. 1995 ) . All subsequent molecular analyses with additional data and sampling recovered a monophyletic family of Burmanniaceae sister to Dioscoreaceae, and therefore part of Dioscoreales (Caddick et al. 2000b(Caddick et al. , 2002aSoltis et al. 2000 ;Davis et al. 2004 ) . ...
This chapter provides a description of all plant families and genera that include putative fully mycoheterotrophic species, excluding initial and partial mycoheterotrophs. The overview covers a total of 17 families, 101 genera, and ca. 880 species. For each family and genus (except for Orchidaceae) a short morphological description is provided followed by notes on taxonomy, distribution, evolution, and ecology. For most genera a line drawing of a representative species is provided. Included families are: Aneuraceae, Burmanniaceae, Corsiaceae, Ericaceae, Gentianaceae, Gleicheniaceae, Iridaceae, Lycopodiaceae, Ophioglossaceae, Orchidaceae, Petrosaviaceae, Podocarpaceae, Polygalaceae, Psilotaceae, Schizaeaceae, Thismiaceae, and Triuridaceae. © 2013 Springer Science+Business Media New York. All rights are reserved.
... Bromeliaceae has been sustained as monophyletic by phylogenetic analyses based on morphological characters ( Stevenson and Loconte 1995 ), as well as molecular ones ( Gaut et al. 1992 ;Duvall et al. 1993 ;Chase et al. 1995 ;Chase et al. 2000 ). Traditionally, three subfamilies are recognized ( Smith and Downs 1979 ): Pitcairnioideae, Tillandsioideae, and Bromelioideae. ...
The genus Quesnelia presently includes 18 species, which occur mainly near the east coast of Brazil from the states of Rio de Janeiro to Bahia. The genus has been divided into two subgenera, Quesnelia and Billbergiopsis. However, its generic and subgeneric delimitation is artificial: in several classifications proposed in the family, different investigators have questioned the naturalness of the group, noting its affinity with species of Aechmea and Billbergia. With the objective of assessing the monophyly of the genus, and evaluating the subgeneric delimitation and the relationship of its species to other genera, a phylogenetic analysis was carried out based on parsimony. The analysis included 33 taxa, with 92 morphological characters. The genera Quesnelia and Aechmea emerged as polyphyletic, and Billbergia as monophyletic. In regard to the subgeneric classification, Quesnelia subgenus Quesnelia emerged as monophyletic, and Quesnelia subgenus Billbergiopsis as polyphyletic. The majority of the species of Quesnelia subgenus Billbergiopsis emerged as the sister group to Billbergia. Even when anatomical and palynological characters were included, the consistency index of the tree obtained was low, indicating high levels of homoplasy. In addition, the majority of clades did not have good statistical support. Therefore, taxonomic changes are not proposed because these would be premature.
... & G.Forst) and Luzuriagaceae (Luzuriaga Ruiz & Pav.). As the publication of molecular systematic studies focused on monocot phylogeny during the past two decades, Smilacaceae have been confirmed to be a member of a monophyletic Liliales, closely related to Philesiaceae, Ripogonaceae and Liliaceae sensu stricto (s.s.) (Chase et al., 1995;Patterson & Givnish, 2002;Fay et al., 2006;Kim et al., 2013;Petersen, Seberg & Davis, 2013). However, the exact position of Smilacaceae in Liliales has been somewhat controversial. ...
Smilacaceae, composed of Smilax and Heterosmilax, are a cosmopolitan family of > 200 species of mostly climbing monocots with alternate leaves characterized by reticulate venation, a pair of petiolar tendrils and usually prickly stems. Although there has been a long history of studying Smilax since Linnaeus named the genus in 1753, the phylogenetic history of this dioecious family remains unclear. Here we present results based on nuclear ribosomal internal transcribed spacer (nrITS) and plastid matK and rpl16 intron DNA sequence data from 125 taxa of Smilacaceae. Our taxon sampling covers all sections of Smilax and Heterosmilax and major distribution zones of the family; species from Ripogonaceae and Philesiaceae are used as outgroups. Our molecular analysis indicates that phylogenetic relationships largely contradict the traditional morphological classification of the family, instead showing a conspicuous geographical pattern among the species clades. The previously recognized genus Heterosmilax was found to be embedded in Smilax. Species in the family are separated into primarily New World and Old World clades, except for a single species lineage, Smilax aspera, that is sister to the remaining species of the family, but with poor statistical support. Ancestral character state reconstructions and examination of distribution patterns among the clades provide important information for future taxonomic revisions and historical biogeography of the group.
... The family was earlier treated as part of Liliaceae sensu lato (subfamily Lilioideae, tribe Scilleae: Engler 1887), but was subsequently regarded as distinct based on data from several sources, including vegetative morphology and anatomy (Fuchsig 1911; Badawi & Elwan 1986), embryology (Schnarf 1929; Wunderlich 1937; Buchner 1948), and seed anatomy (Huber 1969). Following recent molecular phylogenetic analyses, Hyacinthaceae is now assigned to the " higher asparagoid " clade of the order Asparagales, together with other economically and horticulturally important families such as Alliaceae, Amaryllidaceae, Agavaceae, Asparagaceae and Anthericaceae (Chase et al. 1995; Fay & Chase 1996; Fay et al. 2000). The higher asparagoid clade is characterised by the possession of successive microsporogenesis, in contrast to the lower asparagoid grade, in which microsporogenesis is mostly simultaneous (Rudall et al. 1997). ...
As part of an ongoing study of the families of Asparagales for the Anatomy of the Monocotyledons volume series, leaf anatomical characters of a representative sample of 80 species, 32 genera and four subfamilies, including two tribes, of Hyacinthaceae are examined using both light microscopy and scanning electron microscopy. Raised 'pustules' or dark-coloured spots are a feature of leaves of some species. In comparison with many other geophytes, leaves of Hyacinthaceae possess relatively little sclerenchyma. There is a wide range of variation in crystal types in the family. The presence of copious mucilage, probably derived either from laticifer-like idioblasts or 'rhexigenetic' lacunae, is a possible synapomorphy for the subfamily Hyacinthoideae, especially the African tribe Massonieae. Lacunae may have evolved within the family in order to allow rapid leaf expansion in deciduous-leafed species from xeric habitats, although they are entirely absent from Massonieae, which have mostly prostrate leaves with relatively large mesophyll cells and relatively broad epidermal cells. Species with more erect leaves, such as many Hyaantheae, possess thickened periclinal epidermal cell walls, perhaps enabling the relatively narrow leaves to be held in a more upright position. Developmental heterophylly occurs in some genera with erect leaves.
... In his monographic treatment of the Burmanniaceae, Jonker (1938) proposed a subdivision of the family into two tribes: the Burmannieae and the Thismieae. While many present day authors still maintain Jonker's tribal concept (Stevenson & Laconte 1995; Maas – van de Kamer 1998; Kiew 1999; Caddick & al. 2002a; Yang & al. 2002; APG 2003; Sainge & Franke in press) , others have well-founded arguments to follow Agardh (1853), who was the first to consider the Thismieae to represent a distinct family (Chase & al. 1995; Takhtajan 1997; APG 1998; Caddick & al. , 2000b Chase & al. 2000; Caddick & al. 2002b; Neyland 2002; Thiele & Jordan 2002). Since Schlechter (1921), who still distinguished ten genera within the Thismieae, the number of genera was subsequently declining, finally settling down to four (Maas – van de Kamer 1998). ...
Afrothismia saingei, a new species of Burmanniaceae (tribe Thismieae) from Mt. Kupe in South-West Cameroon is described and illustrated. It differs from the other species of the genus Afrothismia by the size of its flower, the long, distinctly heteromorphic tepals with fringed basal extensions, the dorsal invagination of the perianth tube and the unique shape of the internal flange.
... Chanda and Ghosh speculated that these apertures are in fact foursulcate but suggested that thin sections through the aperture were required to show the morphological relationship of the aperture and operculum. The relationships of Dasypogonaceae within the commelinid clade are currently equivocal ( ), although they are sister to Arecaceae in some analyses (Chase et al. 1995). 848 , monosulcate-operculate pollen (SEM). ...
A seco ndary thickening meristem is record ed for the first time in some herbaceous taxa of Asparagales (Herreria montevide nsis and Thysanotus sp inige r), and the new records are assessed in a systematic co ntext. All monocotyl edons lack a vascular cambium, which is typicall y a single per sistent row of cells producing phloem centrifugally and xylem ce ntripetally. However, some monocotyled ons achieve stem thicken ing by means of a different type of later al meristern, either a primary thickening meristem (PTM) near the apex, together with diffuse secondary growth (as in palm s). or a secondary thicken ing meristern (STM) further away from the apex (as in some Asparagales; see Rudall 1991, for review). The PTM and STM are probabl y developmenlally related, although there is complex tissue involvement. In taxa with an STM, the PTM and STM may someti mes be longitudi nally co ntinuous, at least at some stage in the life cycle (Steven son 1980). Virtually all monocotyledons have a PTM , but among tree-forming or woody taxa this has developed along different lines, probabl y more than once, either as an exten sive apical PTM, as in palms, or as an STM, in some Asparagales (see below). The PTM and STM are not homologous with the vascular cambium, as they are tiered (etagen) men stems which produce distinct vascular bundles (of both xylem and phloem) in a parench ymatou s ground tis sue (Fig. 1-3), mainly ce ntripe tally. There are record s of a PTM in some dicotyledon s and other plant groups (DeMaso n 1983), although the hom ology of these requ ires testing . All unequi vocal records of an STM are in the asparagoid clade (sensn Cha se et al. 1995), which compri ses Dahlgren et al.'s (1985) order Asparagales, together with a few membe rs of their Liliales, such as Iridaceae (Rudall 1991). A STM has been re ported in several tree-forming and shru bby asparagoid s: Aga ve, Aloe, Beaucarnea, Calibanus, Cordyiine, Dasylirion, Dra caena, Furcra ea, Klatti a, Nive nia, No lina, Pleomel e, Sansevieria, Witsenia, Xanth orrh oea and Yucca, and also in some more her baceou s taxa with a thick ' woody' rhizome or stem, such as Aphyllanthes, Gasteria.
... According to all information presently available, Xyridaceae (including Abolbodaceae) are the closest relatives to Eriocaulaceae. This is supported by molecular data (Chase et al., 1995;de Andrade et al., 2010) as well as by morphology. Both families have capitula as first-order inflorescences with a strictly acropetal formation of the flowers. ...
Background and AimsInflorescences are thought to be of enormous taxonomic relevance; however, at the same time they are regarded as being notoriously difficult. This is partly due to the conflicting needs of floristics and evolutionary botany, but partly also due to the complicated and confusing terminology introduced by W. Troll and his school.Methods
The branching patterns of representatives of the genera Eriocaulon, Syngonanthus and Paepalanthus have been studied in the field and from preserved material by scanning electron microscopy. Branching patterns and formation sequences have been analysed and documented in longitudinal schemes and diagrams. Repetitive units of different levels are detected and related to the body plans of other species of the family.Key ResultsThe repetition of very few different branching patterns on different levels of complexity may lead to highly complex inflorescences. However, terms are needed only for patterns; levels may be numbered consecutively. While complex inflorescences are often described as additions or aggregations of units, there is some evidence that complex inflorescences are often the result of fractionation of inflorescence meristems.Conclusions
Precise descriptions of inflorescences useful for diagnostics and phylogenetics can be much simpler than they often are today. If complex inflorescences are the result of meristem fractionation, intermediate morphotypes cannot be expected. On the other hand, such intermediate morphotypes should occur if a complex inflorescence is formed following an aggregation pathway. Unless the repetitive patterns shown here are not correlated to complementary gene activities the inflorescences are not fully understood.
Polen de Commelinaceae y Pontederiaceae (Commelinales) de humedales rioplatenses (Argentina). Se estudiaron los caracteres de los granos de polen de Commelinaceae y Pontederiaceae que crecen en áreas de humedales rioplatenses. Los estudios se realizaron con microscopio óptico y electrónico de barrido. Se actualizaron los conocimientos previos sobre elementos de esos grupos, y se describió por primera vez el polen de Pontederia rotundifolia y Tradescantia cerinthoides. Los granos de polen de ambas familias son heteropolares y tienen simetría bilateral. Son monosulcados, equinados en Commelinaceae, y disulcados, con esporodermis lisa, verrugosa o fosulada en Pontederiaceae, este último tipo, nuevo para la familia. La mayoría de los granos presentan membrana del sulco con exina ausente o presente como una delgada capa de nexina. La ornamentación varía en diferentes niveles jerárquicos: permite distinguir tribus y géneros de Commelinaceae, mientras que es un carácter específico en las Pontederiaceae del área.
An analysis of rbcL sequence data for representatives of families of putative sapindalean/rutalean affinity identified a robust clade of core “sapindalean” taxa that is sister to representatives of Malvales. The constitution of this clade approximates the broad concept of Sapindales (sensu Cronquist). Five lineages within the order are recognized: a “rutaceae” clade (Rutaceae, Cneoraceae, Ptaeroxylaceae, Simaroubaceae sensu stricto, and Meliaceae); a “sapindaceae” clade (Sapindaceae, Aceraceae, and Hippocastenaceae); Anacardiaceae plus Burseraceae; Kirkiaceae; and Zygophyllaceae pro parte. Relationships among these groups were only weakly resolved, but there was no support for the recognition of the two more narrowly defined orders, Rutales and Sapindales sensu stricto. Several families that have previously been allied to Sapindales or Rutales show no affinity to the core sapindalean taxa identified with the molecular data, and are excluded from the order: viz. Akaniaceae, Bretschneideraceae, Conneraceae, Coriariaceae, Melianthaceae, Meliosmaceae, Physenaceae, Rhabdodrendraceae, Sabiaceae, Staphyleaceae, Stylobasiaceae, Surianaceae, and Zygophyllaceae sensu stricto.
The tiny seeds, or dust seeds, of heterotrophic plants are usually dispersed by wind. However, most mycoheterotrophic plants grow in the understory of densely vegetated forests, where the wind is less reliable and wind dispersal is likely less efficient. In the present study, we found that seeds of the fully mycoheterotrophic plant Sciaphila secundiflora (Triuridaceae) possess elaiosomes, and that at least one species of ant, Nylanderia flavipes, functions as a dispersal agent of the plant's seeds. This is the first demonstration of myrmecochory in mycoheterotrophic plants, as well as the first report of zoochory in Triuridaceae.
The chloroplast-encoded gene rbcL was sequenced in 30 genera of Commelinaceae to evaluate intergeneric relationships within the family. The Australian Cartonema was consistently placed as sister to the rest of the family. The Commelineae is monophyletic, while the monophyly of Tradescantieae is in question, due to the position of Palisota as sister to all other Tradescantieae plus Commelineae. The phylogenly supports the most recent classification of the family with monophyletic tribes Tradescantieae (minus Palisota) and Commelineae, but is highly incongruent with a morphology-based phylogeny. This incongruence is attributed to convergent evolution of morphological characters associated with pollination strategies, especially those of the androecium and inflorescence. Analysis of the combined data sets produced a phylogeny similar to the rbcL phylogeny. The combined analysis differed from the molecular one, however, in supporting the monophyly of Dichorisandrinae The family appears to have arisen in the Old World, with one or possibly two movements to the New World in the Tradescantieae, and two (or possibly one) subsequent movements back to the Old World; the latter are required to account for the Old World distribution of Coleotrypinae Cyanotinae, which are nested within a New World clade.
The large rbcL analysis published in 1993 (Chase, Soltis, Olmstead et al., 1993) ranks as the largest phylogenetic analysis, molecular or otherwise, ever produced. These data represented over ten years of effort, but significantly the advent of the polymerase chain reaction, PCR, had greatly expanded the numbers of sequences available in the four years just prior to publication. Strategies for production of DNA sequences had become dramatically easier and faster. At the same time that sequencing was becoming more practical, the United States National Science Foundation was approving a large number of molecular systematics proposals, thus making substantial funding available (examine the number of NSF grant numbers listed in the footnotes of the 1993 paper), while Gerard Zurawski (DNAX Corporation) was making available without cost rbcL PCR and internal sequencing primers. Consequently, a large number of laboratories began working on this same plastid locus. To a very large extent and until just recently, protein-coding plastid DNA sequences have been the almost exclusive focus of vascular plant molecular systematists, whereas nearly all published work on other organisms has focused on ribosomal DNA (rDNA; see Chapter 1). By late 1991, the stage had been set for a dramatic increase in the amount of molecular data available on a wide range of seed plants, and this put workers focusing on rbcL in a position to discuss a large-scale analysis. Publication was not necessarily their goal; most simply felt that everyone would benefit from the interaction and that potentially we could use this sort of unpublished, but widely circulated, result as a way to focus better the many individual projects underway.
The recent discovery of diverse fossil flowers and floral organs in Cretaceous strata has revealed astonishing details about the structural and systematic diversity of early angiosperms. Exploring the rich fossil record that has accumulated over the last three decades, this is a unique study of the evolutionary history of flowering plants from their earliest phases in obscurity to their dominance in modern vegetation. The discussion provides comprehensive biological and geological background information, before moving on to summarise the fossil record in detail. Including previously unpublished results based on research into Early and Late Cretaceous fossil floras from Europe and North America, the authors draw on direct palaeontological evidence of the pattern of angiosperm evolution through time. Synthesising palaeobotanical data with information from living plants, this unique book explores the latest research in the field, highlighting connections with phylogenetic systematics, structure and the biology of extant angiosperms.
These comprehensive analyses of systematic relationships within Trilliaceae focused on the relationships within Paris sensu lato (i.e., Paris, Daiswa, and Kinugasa); between species of Trillium; and between Paris sensu lato and Trillium. Seventy species were selected for cladistic analyses and scored for 110 morphological characters; matK and ITS molecular characters were obtained from GenBank for a subset of 26 taxa. Based on the preliminary results, Trillium rivale was used as a functional outgroup. For the subset of 26 species, analysis of the combined ITS and matK sequence data produced six shortest trees; the morphological data, 13 shortest trees; and the combined morphological and molecular data sets, three trees. Analyses of the full morphological data set of 70 species produced 76 shortest trees. Trillium rivale was distinct from both Paris and Trillium and should be placed in its own genus, for which the name Pseudotrillium is proposed. Trillium govanianum was more closely related to Paris than to Trillium but should be retained as a monotypic genus, Trillidium. Trillium and Paris were monophyletic based on molecular as well as morphological evidence. The cladistic analyses strongly support the separation of Paris sensu lato into Daiswa, Kinugasa, and Paris. The monophyly of Trillium after removal of Pseudotrillium and Trillidium was supported in all but the large morphological analysis; subgenus Phyllantherum was monophyletic in all cases, but subgenus Trillium was not monophyletic. Communicating Editor: Jeff H. Rettig
Aquatic, freshwater, rhizomatous perennial. Rhizome dorsiventral, creeping, apparently monopodial, axillary buds sometimes developing into short-stalked bulbils. Leaves distichous or subdistichous, borne on the apex of the rhizome, erect, linear, glabrous not differentiated into petiole and blade, and up to 1 m or more long, triquetrous, submerged or usually emergent, the base somewhat extended and sheathing below; intravaginal scales numerous. Scapes developed on alternate sides of the rhizome, usually separated by an odd number (5, 7, 9) of foliage leaves, erect, terete, glabrous. Inflorescence an umbel-like complex of cymes subtended by (2-)3(-4) bracts. Flowers pedicellate, perfect, protandrous, almost hypogynous, actinomorphic. Perianth of 6 petaloid tepals in 2 whorls, free, white to pink. Stamens 9, the outer whorl with 6 and the inner with 3; filaments flattened; anthers basifixed, 2-thecate, latrorsely and longitudinally dehiscent. Gynoecium of 6 conduplicate carpels arranged in 2 alternate whorls of 3, the carpels connate at the base into a ring, otherwise distinct, each apically narrowed into a crested ventral somewhat decurrent stigma; basally nectariferous on lateral carpel walls; placentation laminar. Ovules numerous, anatropous. Fruit of separate follicles, dehiscing by widening of the ventral sutures. Seeds numerous with ribbed testa and a straight embryo; endosperm lacking.
An account of Velloziaceae species described from cultivated material and well illustrated by colour plates is provided. Colour plates and descriptions of Barbacenia flava and Vellozia intermedia are presented. The original works in which these species were described lack relevant details and illustrations. The descriptions and watercolours presented here provide complementary information for the characterisation of these species.
Fay, M. F. & Chase, M. W.: Resurrection of Themidaceae for the Brodiaea alliance, and recircumscription of Alliaceae, Amaryllidaceae and Agapanthoideae. – Taxon 45: 441‐451. 1996. – ISSN 0040‐0262.
The family Themidaceae Salisb. is resurrected and recircumscribed. It consists of C. 10 genera of cormous plants principally from western N. America (British Columbia to northern Guatemala). These had been included in Alliaceae , as tribe Brodiaeeae (subfamily Allioideae) , but recent molecular and anatomical studies indicate that the group merits familial status. Petronymphe is not a member of this group but rather belongs in Anthericaceae. Agapanthus is misplaced in Alliaceae and, despite its superior ovary, should be considered a member of Amaryllidaceae , as subfamily Agapanthoideae. Alliaceae, Amaryllidaceae , and Agapanthoideae are recircumscribed, and Tulbaghieae raised to subfamilial status.
Medusagyne oppositifolia Baker is the sole member of Medusagynaceae Engl. & Gilg and its phylogenetic position has been unclear. Analysis of rbcL sequence data indicates a close and strongly supported relationship to Ochnaceae and Quiinaceae, but does not resolve the relationships between these taxa. Together the three families form a monophyletic group with a somewhat more distant relationship to other linalean groups including Malpighiaceae, Linaceae and phyllanthoid Euphorbiaceae.
Analyses of morphological and molecular characters for Dioscoreales Hook. f. (Chase & al., 1995b; Caddick & al., 2000a; Caddick & al., 2002) have redefined the order, which now comprises three families, Burmanniaceae, Dioscoreaceae, and Nartheciaceae. Since recent analyses of morphological and molecular data sets (Caddick & al., 2002) have indicated well-supported relationships within Dioscoreaceae R. Br., a formal reclassification of the family is presented here. Dioscoreaceae now contain four distinct genera, Dioscorea, Stenomeris, Tacca (previously in Taccaceae), and Trichopus. The Malagasy endemic Avetra sempervirens is close sister to Trichopus zeylanicus, and is here reclassified as a second species of this genus. The dioecious Dioscoreaceae genera, Borderea, Epipetrum, Nanarepenta, Rajania, Tamus, and Testudinaria, are nested within Dioscorea in phylogenetic analyses (Caddick & al., 2002), and are therefore sunk into it.
Gynoecium and ovule structure was comparatively studied in representatives of the basal monocots, including Acorales (Acoraceae), Alismatales (Araceae, Alismataceae, Aponogetonaceae, Butomaceae, Hydrocharitaceae, JuncÍaginaceae, Limnocharitaceae, Potamogetonaceae, Scheuchzeriaceae, Tofieldiaceae), Dioscoreales (Dioscoreaceae, Taccaceae), and Triuridaceae as a family of uncertain position in monocots. In all taxa studied the carpels or gynoecia are closed at anthesis. This closure is attained in different ways: (1) by secretion without postgenital fusion (Araceae, Hydrocharitaceae); (2) by partly postgenitally fused periphery but with a completely unfused canal (Alismataceae, Aponogetonaceae, Butomaceae, Limnocharitaceae, Scheuchzeriaceae, Dioscoreaceae, Taccaceae); (3) by completely postgenitally fused periphery but with an unfused canal in the centre (Acoraceae, Tofieldiaceae); (4) by complete postgenital fusion and without an (unfused) canal (Juncaginaceae, Potamogetonaceae). In many Alismatales (but without Araceae) carpels have two lateral lobes. The stigmatic surface is restricted to the uppermost part of the ventral slit (if the carpel is plicate); it is never distinctly double-crested (Butomaceae?). Stigmas are commonly unicellular-papillate and secretory in most taxa. The locules are filled with a (often) mucilaginous secretion in a number of taxa. Superficial (probably intrusive) ethereal oil cells were found in the carpel wall of Acorus gramineus (as in Piperales!). Idioblasts in carpels are otherwise rare. A number of basal monocots has orthotropous ovules, which is perhaps the plesiomorphic condition in the group. The presence of almost tenuinucellar (pseudocrassinucellar) ovules is relatively common (Acoraceae, many Araceae, some Alismatales s.s.), whereas completely tenuinucellar ovules are rare and do not characterize larger groups. However, crassinucellar ovules occur in the largest number of families among the study group (basal Araceae, many Alismatales s.s.) The outer integument is always annular in orthotropous ovules. The inner integument is often lobed and it mostly forms the micropyle, whereas the outer integument is always unlobed. Gynoecium structure supports the isolated position of Acoraceae as sister to all other monocots. However, in an overall view, if compared with all other families, Acoraceae clearly shows the greatest similarities with Araceae.
Many molecular studies have shown the monocot order Liliales to be well supported; morphologically, it is defined by synapomorphies of tepalar nectaries and extrorse anthers, in contrast with septal nectaries and introrse anthers commonly found in other monocots, especially Asparagales, with which it was often confused in the past. It comprises c. 1500 species, 67 genera and 9–11 families. Although monophyly is clear, the phylogenetic relationships among some of the families are still unclear. In this study, we examine the inter- and infrafamilial relationships among Liliales in phylogenetic analyses based on four plastid loci (matK, rbcL, atpB and atpF-H). We performed phylogenetic analyses and constructed maximum parsimony and Bayesian trees for 49 genera and 148 taxa in ten families of Liliales sensu Angiosperm Phylogeny Group (APG) III using the combined DNA data. The monophyly of Liliales, except for Corsiaceae (Arachnitis), was strongly supported by both analyses. Campynemataceae were sister to the rest of the order, excluding Corsiaceae. The other families formed two well-defined clades, (Colchicaceae + Alstroemeriaceae) and (Liliaceae, Smilacaceae, (Rhipogonaceae + Philesiaceae)), and one weakly supported clade with Melanthiaceae and Petermanniaceae. Subfamilial and tribal circumscriptions for the three larger families, Colchicaceae, Melanthiaceae and Liliaceae, agreed well with the results of this study, except for the subfamily Calochortoideae of Liliaceae, which was split into two separate clades of Calochortus and Tricyrtis. In addition, we found several taxa with a 10-bp inversion in matK, which could contribute additional homoplasy to these analyses if included without re-coding. Phylogenetic relationships among families of Liliales were better defined here than in a previous molecular analysis, although the placement of Corsiaceae with plastid data remains problematic. Based on these results, reconsideration of the circumscriptions of Rhipogonaceae + Philesiaceae and the subfamilial circumscription for Calochortoideae of Liliaceae is suggested. © 2013 The Linnean Society of London
According to several authors, the Burmanniales are an order that includes the
families Burmanniaceae, Thismiaceae and Corsiaceae. At present, there is no
consensus concerning the circumscription of Burmanniales. All members of
Thismiaceae and Corsiaceae are mycoheterotrophic; Burmanniaceae are composed
of both mycoheterotrophic and autotrophic species. Other than
mycoheterotrophy, there are few characters that unite members of the order.
Because mycoheterotrophy has resulted in extreme vegetative reduction that may
have contributed to homoplasy in the form of character convergence, the
systematics of Burmanniales is problematic. A phylogeny inferred from
large-subunit (26S) ribosome-DNA sequences suggests that the order
Burmanniales is polyphyletic. Specifically, the Burmanniaceae appears as
strongly supported monophyletic group with Thismia as
sister. Corsia is distinct and may be associated with
the Liliales.
Velloziaceae are a monocot family of five genera and c. 250 species (Mello-Silva, 2009). The family is now assigned to the order Pandanales, a small but morphologically diverse monocot order of five families: Cyclanthaceae, Pandanaceae, Stemonaceae (including Pentastemona Steenis), Triuridaceae and Velloziaceae (APG III, 2009). Within Velloziaceae, three genera occur in South America, of which two are endemic to Brazil (Barbacenia Vand., Vellozia Vand.) and the third is Andean (Barbaceniopsis L.B.Sm.). A fourth genus, Xerophyta Juss., grows in tropical Africa and the fifth, Acanthochlamys P.C.Kao, is endemic to China. In this chapter, we present new data on the anther, ovule and embryology of Velloziaceae in the context of ongoing related studies on floral evolution (e.g. Sajo et al., 2010) and the systematics of Pandanales (Furness and Rudall, 2006; Rudall and Bateman, 2006). Earlier studies on ovule and seed development and androecial features were used to delimit the genera of Velloziaceae (Menezes, 1976, 1980). Embryological features of Pandanales are currently poorly known, though some studies suggest that they are potentially systematically useful. For example, Cheah and Stone (1975) reported an unusual megagametophyte in some Pandanus species, in which nucellar nuclei migrate into the developing embryo sac, and tapetum type is unusually variable within Pandanales (Furness and Rudall, 2006).
Morphology and anatomy of a fossil monocotyledon from the late Early Cretaceous and extant monocots are compared. Anatomy was examined based on publications, while leaf morphology, especially the venation, required new observations on fresh and herbarium material. Spixiarum kipea gen. et sp. nov. belongs most likely to Araceae, and may be sister to Orontioideae or is even part of this tribe. Consequently, proto-Araceae were most likely present during the Early Cretaceous in South America. The occurrence of Spixiarum in South America indicates a north Gondwana origin for Orontioideae and and thus may indicate a Gondwanan origin for proto-Araceae. Sedimentological and taphonomic context indicate that Spixiarum had probably a helophytic ecology similar to living Orontioideae and formed possibly the aquatic vegetation of the Crato Lake in association with the Nymphaeales Pluricarpellatia peltata and Jaguariba wiersemana. Early Cretaceous monocotyledon remains have been rarely recorded. It is debatable if their scarceness is a sign of low diversity or may be due to taphonomic/ecologic reasons.
We performed a phylogenetic analysis of 13 families of Asparagales sensu APG III, including 59 Korean taxa representing 10 families based on five plastid regions sequences (matK, rbcL, rpoC1, rps3, and atpF-H intergenic spacer). Parsimony and Bayesian analyses were conducted to clarify the relationships among Korean Asparagales. Among the five regions, matK and atpF-H were the most informative. The most congruent phylogenetic trees were obtained from the combinations of four regions excluding atpF-H. Although both the matK and rbcL gene trees implied that the family system of Asparagales sensu APG III should be modified to include three expanded families, our combined four-regions data set resulted in a highly resolved topology with strong support for the following: (1) Asparagaceae s.l.; (2) Amaryllidaceae s.l.; (3) Hemerocallidoideae of Xanthorrhoeaceae s.l.; (4) Iridaceae; (5) Hypoxidaceae; and (6) Orchidaceae. However, the position of Scilla (Scilloideae of Asparagaceae s.l.) was ambiguous. Hemerocallidoideae was a sister group of both the expanded families of Asparagaceae s.l. and Amaryllidaceae s.l., which are considered the core asparagoids, while Orchidaceae was sister to all other Asparagales. We discuss differentiation of the morphological character within the Korean Asparagales, which clearly suggests that the subfamily Nolinoideae of Asparagaceae s.l. exhibits different character states than the other subfamilies. We compared 10 representative morphological characters to the molecular phylogenetic relationships, and propose that the circumscription within Asparagales sensu APG III needs to be reconsidered.
Despite progress in clarifying the relationships of Dasypogonaceae (four genera, Baxteria, Calectasia, Dasypogon, and Kingia), their infrafamilial relationships and precise affinities within the commelinid clade remain unsatisfactorily resolved. This paper reviews existing data on the systematic affinities of Dasypogonaceae. It also presents new data on floral structure in all four genera, and data on floral ontogeny in Dasypogon. In Dasypogon, Kingia, and Baxteria the ovary is trilocular and septal nectaries are present around the ovary base. In Calectasia, the ovary is unilocular and septal nectaries are entirely absent. Two subfamilial groupings within Dasypogonaceae (Calectasia–Dasypogon and Baxteria–Kingia) are proposed on the basis of leaf anatomy and ovule and ovary morphology. Many floral characters are plesiomorphic in Dasypogonaceae, but some morphological characters support a close relationship with the order Poales sensu lato, especially the epidermal location of the silica bodies. The unusual long-stalked “drumstick” inflorescences of Dasypogon and Kingia resemble those of some Poales, in which flowers are frequently borne on condensed inflorescences. A possible close relationship between Dasypogonaceae and some Poales such as Rapateaceae and Thurniaceae merits further exploration.
The phylogeny of Alismatales remains an area of deep uncertainty, with different arrangements being found in studies that examined various subsets of genes and taxa. Herein we conducted separate and combined analyses of 103 morphological characters and 52 rbcL sequences to explore the controversial phylogenies of the families. Congruence between the two data sets was explored by computing several indices. Morphological data sets contain poor phylogenetic signals. The homology of morphological characters was tested based on the total evidence of phylogeny. The incongruence between DNA and morphological results; the hypothesis of the ‘Cymodoceaceae complex’; the relationships between Najadaceae and Hydrocharitaceae; the intergeneric relationships of Hydrocharitaceae; and the evolutionary convergence of morphological characters were analyzed and discussed.
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