Evolution of novel morphological and reproductive traits in a clade containing Antirrhinum majus (Scrophulariaceae)

Phylogenetic analysis of DNA sequences of the chloroplast genes rcbL and ndhf revealed a highly supported clade composed of the families Plantaginaceae, Callitrichaceae, and Hippuridaceae in close association with the model organism Antirrhinum majus and other members of family Scrophulariaceae. Plantago has miniature actinomorphic wind-pollinated flowers that have evolved from zygomorphic animal-pollinated precursors. The aquatic Hippuridaceae have reduced windpollinated flowers with one reproductive organ per whorl, and three, rather than four, whorls. In monoecious aquatic Callitrichaceae, further reduction has occurred such that there is only one whorl per flower containing a single stamen or carpel. Optimization of character states showed that these families descended from an ancestor similar to Antirrhinum majus. Recent studies of plant developmental genetics have focused on distantly related species. Differences in the molecular mechanisms controlling floral development between model organisms are difficult to interpret due to phylogenetic distance. In order to understand evolutionary changes in floral morphology in terms of their underlying genetic processes, closely related species exhibiting morphological Variation should be examined. Studies of genes that regulate morphogenesis in the clade described here could aid in the elucidation of a general model tot such fundamental issues as how changes in floral symmetry, organ number, and whorl number are achieved, as well as providing insight on the evolution of dicliny and associated changes in pollination syndrome.     

Morphology of Maundia supports its isolated phylogenetic position in the early‐divergent monocot order Alismatales

According to recent molecular phylogenetic data, the rare Australian endemic Maundia triglochinoides does not form a clade with taxa traditionally classified as members of Juncaginaceae. Therefore, views on the morphological evolution and taxonomy of Alismatales require re‐assessment. As the morphology of Maundia is poorly known and some key features have been controversially described in the literature, the flowers, fruits, inflorescence axes and peduncles were studied using light and scanning electron microscopy. Inflorescences are bractless spikes with flowers arranged in trimerous whorls. Except in the inflorescence tip (where the flower groundplan is variable), flowers possess two tepals in transversal‐abaxial positions, six stamens in two trimerous whorls and four carpels in median and transversal positions. Fruits are indehiscent. The shared possession of orthotropous ovules supports the molecular phylogenetic placement of Maundia as sister to a large clade including Potamogetonaceae and related families. Maundia and Aponogeton spp. share the same highly unusual floral groundplan, a homoplastic similarity that can be explained by spatial constraints in developing inflorescences. The nucellar coenocyte of Maundia appears to be unique among monocots. As Maundia exhibits a mosaic of features characteristic of other families of tepaloid core Alismatales, its segregation as a separate family is plausible. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 12–45.     

Anatomical studies on Sinofranchetia chinensis (Lardizabalaceae) and their systematic significance

The anatomical structures of the Chinese endemic and monotypic genus Sinofranchetia (Lardizabalaceae) are described. There are reticulate, simple-reticulate, scalariform, simple-scalariform and simple perforations in vessel elements as well as in the fibres in the secondary wood of the roots and the stems. The node is trilacunar. The vascular bundles in the petiole are arranged in a ring. Clustered crystals occur in the parenchymatous cells of stems, petioles and pedicles. Leaf stomata are actinocytic. The nodes of sepals, petals and stamens both in male and female flowers are unilacunar and one-traced. There are three sterile carpels with two to three traces in the male flowers, three fertile carpels with two to three traces, and sometimes three sterile carpels lacking a vascular supply. In morphology, the anther dehiscence mechanism and pollen in the female flowers are the same as in the male flowers, such that the so-called female flowers might be bisexual in morphology. In comparing morphology, the sex of the flowers and the perforations of the vessel elements in Sinofranchetia with Decaisnea and other genera of the Lardizabalaceae, Sinofranchetia is considered a basic group at least as the same evolutionary level in the family as Decaisnea . © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 271–281.     

Phylogenetic relationships and evolution of Crassulaceae inferred from matK sequence data

Chloroplast gene matK sequence data were used to estimate the phylogeny of 112 species of Crassulaceae sampled from 33 genera and all six recognized subfamilies. Our analyses suggest that five of six subfamilies recognized in the most recent comprehensive classification of the family are not monophyletic. Instead, we recovered a basal split in Crassulaceae between the southern African CRASSULA: clade (Crassuloideae) and the rest of the family (Sedoideae). These results are compatible with recent studies of cpDNA restriction site analyses. Within Sedoideae, four subclades were also recovered: KALANCHOE:, Leucosedum, Acre, and AEONIUM:; evidence also exists for a TELEPHIUM: clade and SEMPERVIVUM: clade. The genus SEDUM: is highly polyphyletic with representatives spread throughout the large Sedoideae clade. Sympetaly and polymerous flowers have arisen multiple times in Crassulaceae and thus are not appropriate characters upon which to base subfamilial limits, as has been done in the past. One floral character, haplostemy, appears to be confined to the well-supported CRASSULA: clade. Our analyses suggest a southern African origin of the family, with subsequent dispersal northward into the Mediterranean region. From there, the family spread to Asia/eastern Europe and northern Europe; two separate lineages of European Crassulaceae subsequently dispersed to North America and underwent substantial diversification. Our analyses also suggest that the original base chromosome number in Crassulaceae is x = 8 and that polyploidy has played an important role in seven clades. Three of these clades are exclusively polyploid (SEMPERVIVUM: clade and two subclades within the KALANCHOE: and AEONIUM: clades), whereas four (Crassula, Telephium, Leucosedum, and ACRE: clades) comprise both diploid and polyploid taxa. Polyploidy is particularly rampant and cytological evolution especially complex in the ACRE: clade.     

Phylogeny, concerted convergence, and phylogenetic niche conservatism in the core Liliales: insights from rbcL and ndhF sequence data

Calochortus and the family Liliaceae s.s. have often been considered each other's closest relatives, based partly on their shared possession of bulbs, visually showy flowers, winged wind-dispersed seeds, and narrow parallel-veined leaves. We present a well-supported molecular phylogeny for these groups and their close relatives in the core Liliales, based on sequence variation in the chloroplast-encoded rbcL and ndhF genes. This analysis identifies Liliaceae s.s. as monophyletic. including one clade (((Lilium, Fritillaris, Nomocharis), Cardiocrinum), Notholirion) that appears to have diversified in the Himalayas roughly 12 million years ago and another ((Erythronium, Tulipa), (Gagea, Lloydia)) that arose in East Asia at about the same time. Medeola and Clintonia are sister to Liliaceae s.s. and bear rhizomes, inconspicuous flowers, fleshy animal-dispersed fruits, and broad reticulate-veined leaves. Calochortus is sister to Tricyrtis; both Tricyrtis and the neighboring clade of Prosartes-Streptopus-Scoliopus share several of the traits seen in Medeola-Clintonia. The core Liliales thus provide compelling examples of both concerted convergence and phylogenetic niche conservatism. Invasion of open, seasonal habitats was accompanied by the independent evolution of bulbs, showy flowers, wind-dispersed seeds, and narrow parallel-veined leaves in Calochortus and Liliaceae s.s. Conversely, persistence in shady habitats was accompanied by the retention of rhizomes, inconspicuous flowers, animal-dispersed seeds, and broad reticulate-veined leaves in their sister groups. We advance arguments for the context-specific adaptive value of each of these traits, as well as evidence of parallel trends in other groups. Concerted convergence--convergence in several different traits, favored by the same shared set of ecological conditions, in two or more lineages--is an important evolutionary process that can mislead evolutionary analyses based solely on phenotypic variation.     

Specialization for hawkmoth and long-proboscid fly pollination in Zaluzianskya section Nycterinia (Scrophulariaceae)

The evolution of pollination systems has been investigated in Zaluzianskya section Nycterinia (Scrophulariaceae), a group characterized by very long-tubed flowers (up to 60 mm) that open either only at night (19 species) or during the day (one species). Field observations of three species: Z. natalensis , Z. elongata and Z. pulvinata , revealed that their flowers are pollinated shortly after dusk by hawkmoths. Further observations showed that Z. microsiphon , the sole species to have flowers that open only during the day, is pollinated exclusively by long-proboscid flies belonging to the family Nemestrinidae. A phylogenetic analysis of Z. sect. Nycterinia based on morphological characters indicates that Z. microsiphon evolved within a clade of night-flowering species. The shift from hawkmoth to long-proboscid fly pollination in Zaluzianskya was probably facilitated because long corolla tubes, white petals and dilute nectar in hawkmoth-pollinated flowers are pre-adaptations for pollination by long-proboscid flies. However, the autapomorphic features of Z. microsiphon , such as zygomorphy, diurnal flower anthesis and lack of scent, can be regarded as adaptations for long-proboscid fly pollination. Breeding system experiments on Z. natalensis and Z. microsiphon revealed the existence of genetic self-incompatibility. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 138 , 17–27.     

Phylogeny and morphological evolution of tribe Menispermeae (Menispermaceae) inferred from chloroplast and nuclear sequences

The Menispermaceae family contains ca. 72 genera with 450 species that are almost entirely tropical. Its phylogeny at the tribal level has never been examined using molecular data. Here we used DNA sequences of the chloroplast matK gene and trnL-F regions, and the nuclear ITS region to study the delimitation and position of the tribe Menispermeae within the family and its subtribal monophyletic groups. Family-wide phylogenetic analyses of the chloroplast data produced two strongly supported clades. The first clade contains two subclades: Coscinieae including Arcangelisia and Anamirta, and Tinosporeae sensu lato including Fibraureae, supported by morphological characters, such as traits of the cotyledon, stylar scar and embryo. The second clade consists of the tribes Menispermeae sensu DC. and Tiliacoreae Miers. All our analyses surprisingly recognized that tribe Menispermeae is not monophyletic unless tribe Tiliacoreae is included, suggesting that characters of cotyledon and stylar scar are very important for the infrafamilial classification, and that endosperm presence vs. absence was over-emphasized in traditionally tribal division of the family. Our topologies indicate a secondary loss of endosperm. The monophyly of two subtribes of the tribe Menispermeae, Stephaniinae and Cissampelinae, is supported by the cpDNA and ITS data, as well as by morphological characters, including aperture types and shapes, and colpal membrane features of pollen grains, and sepal number of male flowers. The Cocculinae was recognized as a paraphyletic group containing the remaining genera of the tribe Menispermeae.     

“金虎尾路线”植物的花进化与传粉转变

以金虎尾科植物地理分布格局及迁移历史总结出来的“金虎尾路线”, 是解释热带植物洲际间断分布与长距离扩散格局的重要模式。金虎尾路线阐明了金虎尾科植物历史时期7次独立的从起源中心(南美洲)向旧世界(非洲和亚洲)的洲际长距离扩散事件。本文总结了金虎尾路线植物起源地与扩散地主要类群的花部特征与传粉系统, 以探讨这些类群及类似植物长距离扩散后的花进化与传粉转变等适应规律。金虎尾科的南美洲类群都有分泌油脂的萼片腺体, 花的形态结构非常保守, 是与美洲当地特有的条蜂科集油蜂长期协同进化的结果。金虎尾科的非洲类群花保守性消失, 花白色、辐射对称且无萼片腺体, 繁育系统为雄花-两性花异株(功能性的雌雄异株), 传粉者是采集花粉的蜜蜂科昆虫。亚洲的一些属发生了类似非洲类群的泛化适应转变, 但风筝果属(Hiptage)出现了镜像花、异型雄蕊和极度反折的花瓣, 且传粉者是亚洲特有的大蜜蜂(Apis dorsata), 显示出了非常特化的适应性转变。风筝果属所在支系的现存类群涵盖了南美洲、中美洲、非洲和亚洲等地的地方特有属, 体现了金虎尾路线整个迁移历史过程, 是认识金虎尾路线及其进化适应规律的关键类群, 值得在今后的研究中加以重视。     

Cladistic Analysis of A Problematic Ammonite Group: the Hamitidae (Cretaceous, Albian–turonian) and Proposals for New Cladistic Terms

The Hamitidae are a family of mid–Cretaceous heteromorph ammonites including lineages leading to four other families. Problems are outlined in trying to describe the phylogeny of completely extinct groups such as these heteromorph ammonites using the existing cladistic terminology, which is largely concerned with extant taxa and their ancestors. To solve these problems, two new terms are proposed: †crown groups and †stem groups, which are equivalent to crown and stem groups in terms of the evolutionary history of a clade, but are not defined on the basis of extant taxa. Instead they are defined by the topology of the phylogenetic tree, the †crown group being a clade defined by synapomorphies but which gave rise to no descendants. A †stem group is a branch of a phylogenetic tree which comprises the immediate sister groups of a given †crown group but is not itself a clade. Examples of these terms are described here with reference to the phylogeny of the Hamitidae and their descendants. The Hamitidae are paraphyletic and form †stem groups to a number of †crown groups, namely the Anisoceratidae, Baculitidae, Scaphitidae, and Turrilitidae. The definitions of the genera and subgenera are refined with respect to the type species and the clades within which they occur, and four new genera are described: Eohamites , Helicohamites , Sziveshamites , and Planohamites .     

Phylogeny of the Cucurbitales based on DNA sequences of nine loci from three genomes: implications for morphological and sexual system evolution

The Cucurbitales are a clade of rosids with a worldwide distribution and a striking heterogeneity in species diversity among its seven family members: the Anisophylleaceae (29-40 species), Begoniaceae (1400 spp.), Coriariaceae (15 spp.), Corynocarpaceae (6 spp.), Cucurbitaceae (800 spp.), Datiscaceae (2 spp.), and Tetramelaceae (2 spp.). Most Cucurbitales have unisexual flowers, and species are monoecious, dioecious, andromonoecious, or androdioecious. To resolve interfamilial relationships within the order and to polarize morphological character evolution, especially of flower sexual systems, we sequenced nine plastids (atpB, matK, ndhF, rbcL, the trnL-F region, and the rpl20-rps12 spacer), nuclear (18S and 26S rDNA), and mitochondrial (nad1 b/c intron) genes (together approximately 12,000 bp) of 26 representatives of the seven families plus eight outgroup taxa from six other orders of the Eurosids I. Cucurbitales are strongly supported as monophyletic and are closest to Fagales, albeit with moderate support; both together are sister to Rosales. The deepest split in the Cucurbitales is that between the Anisophylleaceae and the remaining families; next is a clade of Corynocarpaceae and Coriariaceae, followed by Cucurbitaceae, which are sister to a clade of Begoniaceae, Datiscaceae, and Tetramelaceae. Based on this topology, stipulate leaves, inferior ovaries, parietal placentation, and one-seeded fruits are inferred as ancestral in Cucurbitales; exstipulate leaves, superior ovaries, apical placentation, and many-seeded fruits evolved within the order. Bisexual flowers are reconstructed as ancestral, but dioecy appears to have evolved already in the common ancestor of Begoniaceae, Cucurbitaceae, Datiscaceae, and Tetramelaceae, and then to have been lost repeatedly in Begoniaceae and Cucurbitaceae. Both instances of androdioecy (Datisca glomerata and Schizopepon bryoniifolius) evolved from dioecious ancestors, corroborating recent hypotheses about androdioecy often evolving from dioecy.     

Reversal versus specialization in floral morphological evolution in Petrocosmea (Gesneriaceae)

A widely held hypothesis in evolution is that adaptive specialization constrains the potential direction of future evolutionary change and thus may be irreversible, also known as Dollo's law. However, this hypothesis has long been subject to debate in evolutionary biology. Floral specialization is intriguing as it is usually linked to reproductive isolation and could affect speciation. Here, following the discovery of four new taxa, we observed some interesting phenomena of reversal versus specialization in morphology in a clade with the most specialized flowers in the genus Petrocosmea. In the phylogenetic tree based on sequences of multiple DNA regions, the morphological reversals, especially the regain of a long corolla tube, are nested within the branches characteristic of normally specialized flowers with a short corolla tube and highly specialized zygomorphy. Our results indicate that the highly specialized floral organ of this clade is still actively evolving in multiple branches toward specialization while reversals to different ancestral states occur in some branches. Great disturbance of ecological environment is likely a crucial factor affecting trait reversibility, such as the rapid uplift of the Himalayan–Tibetan plateau. The four new taxa are treated herein taxonomically. The flowers of this clade represent an interesting model to explore the genetic basis underlying the evolutionary reversal versus specialization and the interplay between genetic factors and environmental variables.     

Developmental morphology of female flowers of Gyrostemon and Tersonia and floral evolution among Gyrostemonaceae

Floral simplifications and specializations in the evolution of Gyrostemonaceae have confused the systematics of the family. Recent phylogenetic analyses have demonstrated their placement among Capparales. This investigation presents a phylogenetic analysis of Gyrostemonaceae, demonstrating that Codonocarpus and Gyrostemon form a clade that is the sister group of Cypselocarpus, Tersonia, and Walteranthus. These phylogenetic results and data on development of Gyrostemon and Tersonia are used to discuss the morphology and evolutionary diversification of female flowers of Gyrostemonaceae. The uniseriate perianth of Gyrostemonaceae consists of four to eight tepals with an unusual lateral to median developmental sequence. The female flowers of Gyrostemon and Tersonia display no distinctive evidence of an androecium, although the former has late-forming, primordium-like structures positioned between the tepals and gynoecium that may be the vestiges of either a second perianth series or the androecium. The gynoecium of Gyrostemonaceae is syncarpous, although the two main clades in the family differ in the expression of ovarian synorganization. The Codonocarpus–Gyrostemon clade is unusual in having largely separate carpels that are only syncarpous because the ventral side of each is formed by the flank of the floral apex. All Gyrostemonaceae, however, incorporate the flank of the floral apex as the ventral side of the carpel, and this is the location of ovule development. On the basis of its placement in a clade that includes Tersonia and Walteranthus, the uniloculate and uniovulate gynoecium of Cypselocarpus may be pseudomonomerous. All Gyrostemonaceae have large stigmas that are typical of anemophilous taxa, and they differ from most other Capparales in this attribute. Among Capparales, Gyrostemonaceae may be most similar to Ochradenus (Resedaceae), which also appears to be anemophilous. It is unclear whether the similarities of Ochradenus and Gyrostemonaceae are homologies, indicative of a close relationship between the two groups, or evolutionary parallelisms associated with separate shifts to anemophily.     

Evolution of reproductive traits in the mahagony family (Meliaceae)

Meliaceae are a mostly pantropical family in the Sapindales, bearing flowers typically provided with a staminal tube, formed by filaments that are fused partially or totally. Nevertheless, several genera of subfamily Cedreloideae have free stamens, which may be adnate to an androgynophore in some taxa. The fact that the family exhibits a wide diversity of floral and fruit features, as well as of sexual systems and pollination syndromes, presents interesting questions on the evolutionary processes that might have taken place during its history. In this study, we analyzed the distribution of 20 reproductive morphological traits of Meliaceae, upon an available molecular phylogenetic framework, using 31 terminals from the family's two main clades (Cedreloideae and Melioideae), plus six Simaroubaceae taxa as outgroup. We aimed to identify and/or confirm synapomorphies for clades within the family and to develop hypotheses on floral evolution and sexual systems in the group. Our reconstruction suggests that the ancestor of Meliaceae was possibly provided with united stamens and unisexual flowers in dioecious individuals, with a subsequent change to free stamens and monoecy in the ancestor of Cedreloideae. Most characters studied show some degree of homoplasy, but some are unique synapomorphies of clades, such as the haplostemonous androecium. An androgynophore defines the Cedrela‐Toona clade. The comparative approach of our study and the evolutionary hypotheses generated herein reveal several aspects demanding further structural investigation, and possible evolutionary pathways of the reproductive structures along with the lineages' diversification, mostly related to the specialization of sexual systems, floral biology, and dispersal strategies.     

A new genus and species of fimbriid bivalve from the Kimmeridgian of the western Pontides, Turkey, and the phylogeny of the Jurassic Fimbriidae

A new lucinoidean genus and species, Cerkesia contiae gen. et sp. nov., from the Kimmeridgian carbonate platform deposits of the Çerkes area (western Pontides, Turkey), is described and placed in the family Fimbriidae. Morphological comparisons reveal similarities with Fimbria , Cyclopellatia and Sphaera . A phylogenetic analysis supports the monophyly of the family. Similarities between Cerkesia and Fimbria determine inclusion of these genera in the same clade. The unusual features characterizing Cerkesia closely relate it to Cyclopellatia . Sphaera belongs to a different clade and its similarities with Cerkesia are owing to convergent and plesiomorphic characters. Cerkesia was a slow and shallow burrower. The morphology of the anterior adductor scar suggests a mode of life comparable to that of Fimbria but other features indicate adaptive strategies typical of the Lucinidae. The associated fossils and facies reveal that Cerkesia lived in soft and sandy substrates of quiet and protected areas of the carbonate platform under stressful environmental conditions. The appearance of Cerkesia in the western Pontides marks the eastward expansion of a clade that developed principally in the European seas during the Late Jurassic while the Sphaeriola-Sphaera clade was confined to the southernmost Tethyan margin.     

Patterns of diversification in New Zealand Stylidiaceae

Phylogenetic analysis of ITS and rbcL sequences show that New Zealand Stylidiaceae fall into two distinct lineages differing in species richness. Each lineage represents a unique dispersal event to New Zealand occurring at different times during the evolutionary history of the family. One lineage comprises seven species of Forstera and Phyllachne, while the other consists solely of Oreostylidium subulatum. The origin of the Forstera/Phyllachne lineage in New Zealand is equivocal; either a South American or a Tasmanian origin is equally parsimonious. Possible sister groups are F. bellidifolia in Tasmania and P. uliginosa in South America. Oreostylidium subulatum has an Australian origin. In our analyses O. subulatum is nested in a clade composed entirely of species of Stylidium, almost all of which are endemic to Australia. Species of Phyllachne share a cushion habit with the outgroup Donatia (Donatiaceae) that may have preadapted them to alpine environments in New Zealand. The New Zealand Stylidiaceae have small, white, actinomorphic flowers that are well adapted to the unspecialized pollinator fauna. Forstera and Phyllachne share this trait with Donatia; however, the small, white flowers of Oreostylidium are a dramatic departure from the colorful, highly specialized flowers of Stylidium.     

Right-left symmetry preservation by flowers of malvaceae family

The flowers of malvaceae family preserves the symmetry between right and left in a peculiar manner. Plots belonging to this family bear two kinds of flowers, right-handed flowers with anticlockwise twisted petals and left-handed flowers with clockwise twisted petals. The branches of the plant prefers production of one type of flowers in excess of the other. There are two distinct types of branches, dextral branches and sinistral branches. Dextral (sinistral) branches produce more right-handed (left-handed) flowers than left-handed (right-handed) flowers. The average percentage of right-handed flowers in a dextral branch is same as that of left-handed flowers in a sinistral branch.     

Morphology and anatomy of the flower of Meliosma (Sabiaceae): implications for pollination biology

Summary The structure and anatomy of mature flowers of four species of Meliosma is investigated using scanning electron and light microscopy. The vasculature of the flower, including the structure of the gynoecium, is described in detail. The mechanism of stamen maturation and pollen release is illustrated and discussed. The existence of an explosive pollination mechanism is questioned for at least part of the species. Flowers are proterandrous and fertile stamens are kept spatially separate from the style by a ring of large staminodes. Anthers are disporangiate by the loss of the adaxial pollen sacs. During maturation the filament bends progressively outwards and releases the pollen on the extension of the connective that acts as a secondary pollen presentation system. The nectary has five appendages topped with stomata secreting abundant nectar. The relationships of Sabiaceae are discussed relative to other early diverging eudicots. The significance of Sabiaceae as an isolated clade is highlighted, although some features point to a link with Menispermaceae.