PALYNOLOGY AND SYSTEMATICS OF CUPHEA (LYTHRACEAE). I. MORPHOLOGY AND ULTRASTRUCTURE OF THE POLLEN WALL

A survey of pollen from 153 species of Cuphea has revealed a remarkable array of morphological forms. The survey involved light and electron microscope investigations of C. crassiflora, jorullensis, and koehneana to determine details of exine morphology, and a more general study of pollen from an additional 150 species. Comparison of pollen types within a single morphological category and within taxonomic groups (i.e., sections or subsections) indicates considerable variation at subgeneric levels. The genus is distinctly eurypalynous, and the extent to which pollen varies among the sections, subsections, species and varieties is probably exceeded by few genera of comparable size. The pollen is also variable within each taxon, but size studies of single-anther lactic acid preparations demonstrate the genus is not polymorphic, as in Lythrum, where pollen polymorphism is associated with heterostyly. The latter phenomenon is as yet unknown in Cuphea. These results reveal that pollen morphology constitutes an important and useful character for taxonomic studies of Cuphea.     

Major clades and a revised classification of Magnolia and Magnoliaceae based on whole plastid genome sequences via genome skimming

With more than 300 species, the Magnoliaceae family represents a major Magnoliid lineage that is disjunctly distributed in Asia and the New World. The classification of Magnolia s.l. has been highly controversial among taxonomists, varying from one genus with several subgenera, sections, and subsections to several (up to 16) genera. We conducted a comprehensive phylogenetic study of Magnoliaceae on the basis of sequences of the complete chloroplast genomes with a broad taxon sampling of 86 species. The phylogenetic analyses strongly support 15 major clades within Magnolia s.l. due to the non‐monophyly of subgen. Magnolia, the previous subgeneric treatment that recognizes three subgenera, is not supported. Based on the phylogenetic, morphological, and geographic evidence, we recognize two subfamilies in Magnoliaceae: Liriodendroideae and Magnolioideae, each with one genus, Liriodendron and Magnolia, respectively. Magnolia is herein classified into 15 sections: sects. Magnolia, Manglietia, Michelia, Gwillimia, Gynopodium, Kmeria, Maingola, Oyama, Rytidospermum, Splendentes, Talauma, Tuliparia, Macrophylla, Tulipastrum, and Yulania.     

HYBRIDIZATION AND CYTOTAXONOMY OF DICENTRA

Biosystematic relationships among species assigned to three subgenera of the genus Dicentra were investigated with respect to hybridization and chromosomal constitution and fertility of the hybrids. Four species of subgenus Dicentra, D. formosa, D. eximia, D. nevadensis, and D. peregrina, were intercrossed in various ways to form diploid, triploid, and tetraploid hybrids. Hybrids at the tetraploid level in this subgenus invariably were highly fertile. Triploid hybrids, as expected, were mostly very sterile. Diploid hybrids varied in this respect, but none was highly fertile. Crosses with two of the remaining four species of subgenus Dicentra produced no hybrids, but abundant seed was obtained in one instance. The two species of the subgenus Chrysocapnos, D. chrysaniha and D. ochroleuca, cross to produce a partially fertile tetraploid hybrid, but cross-pollinations involving these species with those of other subgenera failed. The single species of subgenus Hedycapnos, D. spectablis (diploid) produced no hybrids when cross-pollinated with members of the other two sections. These results are fully concordant with presumed affinities based on morphological similarity In addition, preliminary results of hybridization between the monocotyledonous D. peregrina and a number of dicotyledonous species of Dicentra are reported.     

Age estimates of Frullania (Frullaniaceae,Porellales) main lineages: another example of rapid and recent diversification in liverwort evolution

Frullania Raddi is an extant genus of liverworts (Bryophytes) widespread around the world. It belongs to the family Frullaniaceae Lorch., with a large number of species distributed into several subgenera, sections and subsections according with different morphological classifications. As shown in previous studies, most Frullania sub-generic classifications are supported by molecular data, indicating that morphological characters appear well suitable to discriminate species. However, deep among-clade relationships remain unclear, probably due to the rapid diversification of main clades, paralleling other plant lineages. In this study, we reconstruct Frullania phylogeny based on available molecular data used in previous studies, and we present time estimates for the origin of its main branches. Results supported the monophyly of most subgenera as demonstrated in previous studies and supported a rapid diversification of these main lineages. Time estimates under a relaxed molecular clock and with integrated fossil evidence and nucleotide mutation rates further suggested a Jurassic origin of the genus and a rapid diversification during Palaeogene and Neogene. This may have been influenced by geographical and climate changes during these periods as predicted for most leafy liverworts.     

FLAVONOID EVOLUTION IN DENDROSERIS (COMPOSITAE,LACTUCEAE) FROM THE JUAN FERNANDEZ ISLANDS,CHILE

Leaf flavonoid chemistry was examined from the three subgenera and 11 species of the endemic genus Dendroseris (Compositae, Lactuceae) of the Juan Fernandez Islands, Chile. Eight of the species are restricted to the older island (Masatierra, ca. 4 million years old), which is also closer to the mainland. Three species, one from each subgenus, are restricted to Masafuera, which is younger geologically (1–2 million years old) and 145 km further west of Masatierra. A total of 16 compounds was identified, with the 7-0-glucosides of the flavones apigenin and luteolin accounting for 12 of the constituents. Two glucosides of the flavonol quercetin were detected. Despite considerable interpopulation variation within species, six of the taxa have distinctive flavonoid profiles. Although there are few absolute differences among the subgenera, they can be distinguished chemically. Subgenus Rea contains the greatest number of compounds, and a previous cladistic analysis based on morphological features suggested this subgenus as most primitive. Subgenus Phoenicoseris is considered highly derived morphologically, and it has a reduced flavonoid chemistry. Very little reduction in flavonoid diversity was seen in the morphologically specialized subg. Dendroseris as compared to subg. Rea. A trend in reduction of numbers of compounds was seen for two of the three species on the younger island of Masafuera when compared to their presumed ancestors on Masatierra. Flavonoids of selected species of Hieracium and Hypochaeris, presumptive mainland progenitors of Dendroseris, reveal a close chemical affinity with the former genus.     

Chloroplast DNA restriction analysis and the infrageneric grouping ofAllium (Alliaceae)

The utility of chloroplast DNA variation for checking a recently proposed infrageneric classification of the genusAllium was tested. cpDNA restriction patterns of 49 species representing the main subgenera, sections, and subsections of the existing classification were compared. 363 different fragments generated by 4 restriction enzymes were identified and analysed by UPGMA clustering. The resulting phenogram largely confirms the subgeneric classification based on an integration of morphological and other methods.     

A classification of the New World species of Aster (Asteraceae)

In order effectively to list and comment on the results of cytological investigations inAster in a companion paper, a scheme of infrageneric classification is presented which utilizes the basic chromosome number as a pivotal diagnostic character. Reasons are stated as to why, with the exception ofUnamia Greene which is transferred toSolidago, and the commonly recognized generaLeucelene Greene,Machaeranthera Nees andXylorhiza Nutt., none of the segregate genera previously proposed or recorded in the literature is upheld. Instead, these taxa are being given subgeneric or sectional rank. Two additional subgenera are established to accommodate the species groups traditionally placed in “Aster proper,” which are characterized by having basic chromosome numbers ofx = 5 andx = 8, respectively. Altogether ten subgenera of the genusAster, five of them subdivided further into a total of 24 sections, are recognized as having representative species in the New World. All basionyms and type species are listed, and a number of new combinations and status changes are validated in accordance with the International Rules of Botanical Nomenclature. Where known (from literature and personal research), chromosome numbers are recorded for the species.     

TYPES OF LATICIFERS AND CRYSTALS IN JATROPHA AND THEIR TAXONOMIC IMPLICATIONS

Anatomical examination of 37 species of Jatropha (Euphorbiaceae) has revealed the occurrence of two distinct types of laticifers—articulated, and nonarticulated—in addition to idioblasts that are irregularly shaped individual cells. With the notable exception of J. augustii Pax & K. Hoffm. (section Peltatae (Pax) Dehgan & Webster), these idioblasts occur in the two most advanced sections of the subgenus Curcas (Adans.) Pax, namely Loureira (Cav.) Muell. Arg. ex Pax and Mozinna (Ortega) Pax. The presence of two laticifer types and “idioblastic laticifers” in the genus, in association with the morphological reduction series found with evolutionary advancement are, therefore, significant in the delimitation of sections and subsections. Further evidence is presented by the occurrence of “chambered crystalliferous cells” in subgenus Curcas, but not in subgenus Jatropha (=Adenorhopium Griseb.). Reexamination of laticifers in other genera of the Euphorbiaceae is suggested as a possible means of alleviating the long-standing taxonomic dilemma of this large and morphologically diverse family.     

SYSTEMATIC STUDIES IN THE LEAF ANATOMY OF PALM GENUS SYAGRUS

A systematic survey of the leaf anatomy of 51 taxa in the palm genus Syagrus was made. Cross sections of the middle pinnae of each taxon were embedded, cut, stained, and mounted on slides. Criteria used to distinguish species were based mainly on differences in the laminar surface and hypodermis: frequency and location of various sized veins; frequency, shape, and location of nonvascular fibers; size and shape of midrib and main vascular bundle; and size and shape of expansion cell tissue. The included key to the species employs mostly these characteristics. Each taxon is listed under subgenera and sections in the order presented in a previous article, where gross morphology was used as the basis of classification. Relationships between species of each subgenus and section are discussed in terms of the original arrangement of each group and needed changes. It is significant that certain evolutionary sequences have become unfolded which were not apparent by use of morphological characters alone. Although all relationships have not been completely worked out, this survey has resulted in a revision of my original classification of the genus Syagrus and at the same time has also corroborated certain alignments of species previously based on gross morphology.     

中国菊科一新种——香紫菀北大核心CSCD

该文描述了在中国福建北部山区茫荡山发现的菊科(Compositae)紫菀属(Aster L.)一新种——香紫菀(A.artemisiaodorum X.X.Su,S.P.Chen&L.Ma sp.nov.)。基于2个核基因(ITS、ETS)和1个叶绿体基因(trn L-F)片段构建的系统发育树显示,香紫菀具有独立性,与白舌紫菀[A.baccharoides(Benth.)Steetz]互为姐妹类群,但该种形态与白舌紫菀相差较大,其全部叶两面无毛、总苞筒状、头状花序1.0~1.8 cm、舌状花5~8朵以及全株具芳香气味等特征。形态特征分析与系统发育分析均支持该新种成立。模式标本现存于福建农林大学标本馆。     

An updated classification for the hyper-diverse genus Corydalis (Papaveraceae: Fumarioideae) based on phylogenomic and morphological evidence

The genus Corydalis, with ca. 530 species, has long been considered taxonomically challenging because of its great variability. Previous molecular analyses, based on a few molecular markers and incomplete taxonomic sampling, were clearly inadequate to delimit sections and subgenera. We have performed phylogenetic analyses of Corydalis and related taxa, using 65 shared protein-coding plastid genes from 313 accessions (including 280 samples of ca. 226 species of Corydalis) and 152 universal low-copy nuclear genes from 296 accessions (including 271 samples of Corydalis) covering all 42 previously recognized sections and five independent “series”. Phylogenetic trees were inferred using Bayesian Inference and Maximum Likelihood. Eight selected morphological characters were estimated using ancestral state reconstructions. Results include: (i) of the three subgenera of Corydalis, two are fully supported by both the plastid and nuclear data; the third, subg. Cremnocapnos, is weakly supported by plastid DNA only, whereas in the nuclear data the two included sections form successive outgroups to the rest of the genus; (ii) among all 42 sections and five “series”, 25 sections and one “series” are resolved as monophyletic in both data sets; (iii) the common ancestor of Corydalis is likely to be a perennial plant with a taproot, yellow flowers with a short saccate spur, linear fruits with recurved fruiting pedicels, and seeds with elaiosomes; (iv) we provide a new classification of Corydalis with four subgenera (of which subg. Bipapillatae is here newly described), 39 sections, 16 of which are consistent with the previous classification, 16 sections have been recircumscribed, one section has been reinstated and six new sections are established. Characters associated with lifespan, underground structures, floral spur, fruit and elaiosomes are important for the recognition of subgenera and sections. These new phylogenetic analyses combined with ancestral character reconstructions uncovered previously unrecognized relationships, and greatly improved our understanding of the evolution of the genus.     

Beiträge zur Kenntnis der Sippenstruktur des Genus Parabarium Pierre ex Spire (Apocynaceae)

The genus Parabarium is clearly separated from the other genera of the subtribe Urceolinae by different characters. The inflorescences of their taxa are heterocladical pleiothyrses, trunk synflorescences, or pleiochasiums. Further morphological and anatomical investigations deals with flower buds, leaves, pollen grains, fruits, seed, and other parts of the plants. Basing on the character analysis resulting from this the genus Parabarium is divided into two subgenera with two sections and two subsections enclosing 16 species. Two of the latter are described as news, further two varieties are set up as new ones. Two species are transferred from the genus Ecdysanthera into the genus Parabarium. Further three Parabarium species are inserted, and some more taxonomical corrections are made. — The possible relations within the genus Parabarium are made. Based on these results P. micranthum and therefor the subsection Parabarium (subgenus Parabarium) have to be regarded as the more original ones in face of P. quintaretii and the subgenus Pleiochasium. The section Burmanicum of the subgenus Parabarium has an exceptional position within the genus     

Phylogeny of mosquitoes of tribe Culicini (Diptera: Culicidae) based on morphological diversity

Harbach, R. E., Kitching, I. J., Culverwell, C. L., Dubois, J. & Linton, Y.‐M. (2012). Phylogeny of mosquitoes of tribe Culicini (Diptera: Culicidae) based on morphological diversity. —Zoologica Scripta, 41, 499–514. Relationships among taxa of the mosquito tribe Culicini are explored using 169 morphological characters from 86 exemplar species representing the four genera and 26 subgenera of Culicini, most species groups and subgroups of subgenus Culex and an outgroup of five species from five other tribes. We analysed the data set, with multistate characters treated as unordered, under implied weights with values of K ranging from 1 to 20, implemented by TNT. Each analysis, except K = 4, produced a single most parsimonious (fittest) cladogram (MPC). The topology of the ingroup was identical for K = 6–11, whereas the MPCs for K = 14–20 differed only in the position of a single species, which occupied the same position in the K = 16 and K = 6–11 topologies. The K = 9 and K = 16 trees were given further consideration. In both these cladograms, Lutzia is sister to a clade comprising genera Culex, Deinocerites and Galindomyia. The two topologies have 13 clades in common, but their arrangements differ primarily because of Culex (Culex) duttoni acting as a ‘rogue’ taxon. We evaluated the effect of removing this species from the K = 9 and K = 16 analyses and chose the refined K = 9 topology as the best hypothesis of relationships within Culicini. Genus Culex is not monophyletic because it includes Deinocerites and Galindomyia as derived members of the New World subgenera. With the exception of subgenera Culex, Eumelanomyia and Neoculex, there is strong support for the monophyly of all genera and subgenera. Subgenus Culex would be monophyletic were four other subgenera included and three other taxa (the Afrotropical Cx. duttoni, Neotropical Cx. apicinus and the Australasian Atriceps Group) excluded.     

A new infrageneric classification and phylogenetic trends in the genusRhododendron (Ericaceae)

Recent studies have improved the infrageneric classification ofRhododendron, including my own investigations on flavonoids and anthocyanins as chemosystematic markers. From a synoptical comparison of morphological, anatomical and phytochemical characters a new system for the genus is proposed. Phylogenetic character progressions and relationships among subgenera, sections and subsections are discussed and illustrated. Key positions for subg.Candidastrum between chori subgenerumRhododendron andNomazalea, and for subg.Choniastrum between chori subgenerumHymenanthes andNomazalea are suggested.     

MOLECULAR PHYLOGENY OF THE HETEROTROPHIC DINOFLAGELLATES,PROTOPERIDINIUM, DIPLOPSALIS AND PREPERIDINIUM (DINOPHYCEAE), INFERRED FROM LARGE SUBUNIT rDNA1

The genera Protoperidinium Bergh, Diplopsalis Bergh, and Preperidinium Mangin, comprised of species of marine, thecate, heterotrophic dinoflagellates in the family Protoperidinaceae Balech, have had a confused taxonomic history. To elucidate the validity of morphological groupings within the Protoperidinium and diplopsalids, and to determine the evolutionary relationships between these and other dinoflagellates, we undertook a study of molecular phylogeny using the D1–D3 domains of the large subunit (LSU) of the rDNA. Based on morphology, the 10 Protoperidinium species examined belonged to three subgenera and five morphological sections. Two diplopsalid species were also included. Single‐cell PCR, cloning, and sequencing revealed a high degree of intraindividual sequence variability in the LSU rDNA. The genus Protoperidinium appeared to be recently divergent in all phylogenetic analyses. In maximum parsimony and neighbor joining analyses, Protoperidinium formed a monophyletic group, evolving from diplopsalid dinoflagellates. In maximum likelihood and Bayesian analyses, however, Protoperidinium was polyphyletic, as the lenticular, diplopsalid heterotroph, Diplopsalis lenticula Bergh, was inserted within the Protoperidinium clade as basal to Protoperidinium excentricum (Paulsen) Balech, and Preperidinium meunieri (Pavillard) Elbrächter fell within a separate clade as a sister to the Oceanica and Protoperidinium steidingerae Balech. In all analyses, the Protoperidinium were divided into two major clades, with members in the Oceanica group and subgenus Testeria in one clade, and the Excentrica, Conica, Pellucida, Pyriforme and Divergens sections in the other clade. The LSU rDNA molecular phylogeny supported the historical morphologically determined sections, but not a simple morphology based model of evolution based on thecal plate shape.     

Karyological and molecular characterisation of subgenus Vicia (Fabaceae)

In the present report, we have analysed the subgenus Vicia by karyological and molecular approaches with the aim to clarify the relationships among Vicia species included in this subgenus by previously evidenced morphological investigations. Multivariate analysis using several karyomorphological parameters in addition to symmetry indices has allowed the construction of a dendrogram of linkage distances very useful to compare and to include in a phylogenetic tree obtained by internal transcribed spacer (ITS) rDNA sequences. Moreover, a separate analysis was performed combining our molecular data on ITS sequences with those reported in the literature for the section Vicilla. Our analyses partly confirm the monophyletic status of the various sections in which the subgenus Vicia has been divided, however questioning, in some cases, the real need to maintain all the nine sections so far accepted and the placement of some individual species in the two subgenera Vicia and Vicilla.     

EXPERIMENTAL ASSESSMENT OF REPRODUCTIVE INTERACTIONS BETWEEN SYMPATRIC ASTER AND ERIGERON (ASTERACEAE) IN INTERIOR ALASKA

Erigeron glabellus and Aster sibiricus have similar flowers, share pollinators, but bloom sequentially in interior Alaska. Both species depend on insect pollination for seed set: the Erigeron is self-incompatible, and the Aster is apparently self-compatible but allogamous. To test the hypothesis that sequential blooming is maintained by natural selection generated by reproductive interference, we manipulated the flowering time of Erigeron, forcing it to bloom simultaneously with Aster, and measured female fecundity in both species. We found no evidence of reduced female fecundity in either species caused by the presence of the sympatric “competitor” or by artificial pollination with the heterospecific pollen prior to conspecific pollination. Two-species mixtures of simultaneously blooming Aster and Erigeron experienced significant interspecific visitation, which may, under natural conditions, cause loss of pollen to alien stigmas and depressed male fecundity, at least in Erigeron. We found no evidence that sequential blooming in Erigeron and Aster is maintained by depressed female fecundity through pollinator sharing. If sequential blooming is maintained by natural selection, it seems more likely to be the result of selection generated by depressed male fitness through pollen loss to alien stigmas.     

Phylogenetic analysis based on structural and combined analyses of Rhus s.s. (Anacardiaceae)

Structural data were combined with trnL‐F and internal transcribed spacer sequences from other studies and with new sequences representing ten additional species to clarify the phylogenetic relationships of Rhus s.s. These data indicate that Rhus s.s and both subgenera, Rhus and Lobadium, are monophyletic. The genus Rhus is supported as monophyletic by the presence of red glandular hairs on the berries and inflorescence axis, cilia on the sepals and glands on the leaf blades. Subgenus Rhus can be identified by the presence of more than seven resin channels in the petiole, weakly percurrent tertiary veins and a type I vascular system in the mid‐vein. Subgenus Lobadium is characterized by the presence of short bracteoles and pedicels. This subgenus is divided into four sections, Lobadium, Rhoeidium, Styphonia and Terebinthifolia. Section Lobadium has trifoliate leaves; section Rhoeidium is monotypic, including only Rhus microphylla; section Styphonia is supported by five synapomorphies, including an incomplete marginal vein, fibres in the petiole, a thick cuticle, two layers of palisade parenchyma and prismatic crystals in the mesophyll; and section Terebinthifolia has gelatinous xylary fibres in the petiole. Hypotheses about the evolutionary changes of these characters are presented based on the cladograms. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176 , 452–468.     

New insights into the phylogenetics and biogeography of Arum (Araceae): unravelling its evolutionary history

The heat‐ and odour‐producing genus Arum (Araceae) has interested scientists for centuries. This long‐term interest has allowed a deep knowledge of some complex processes, such as the physiology and dynamics of its characteristic lure‐and‐trap pollination system, to be built up. However, mainly because of its large distributional range and high degree of morphological variation, species' limits and relationships are still under discussion. Today, the genus comprises 28 species subdivided into two subgenera, two sections and six subsections. In this study, the phylogeny of the genus is inferred on the basis of four plastid regions, and the evolution of several morphological characters is investigated. Our phylogenetic hypothesis is not in agreement with the current infrageneric classification of the genus and challenges the monophyly of several species. This demonstrates the need for a new infrageneric classification based on characters reflecting the evolution of this enigmatic genus. To investigate the biogeography of Arum deeply, further spatiotemporal analyses were performed, addressing the importance of the Mediterranean basin in the diversification of Arum. Our results suggest that its centre of origin was the European–Aegean region, and that major diversification happened during the last 10 Myr. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163 , 14–32.