Quincuncial Aestivation Research Articles

Comparative floral development reveals novel aspects of structure and diversity of flowers in Cannabaceae

AbstractSpecies of Cannabaceae are wind pollinated, have inconspicuous and reduced flowers that are pistillate, staminate and apparently perfect on the same individual or on different individuals, with a single-whorled perianth and a pseudomonomerous gynoecium. Our objective is to understand the developmental processes that lead to such a reduced flower morphology and polygamy in Cannabis sativa, Celtis iguanaea and Trema micrantha. Floral buds and flowers were processed for surface, histological examinations and 3D reconstructions of vasculature. The single-whorled perianth is interpreted as a calyx because the organs are robust, have a broad base, an acute apex and quincuncial aestivation and are opposite the stamens. Petals are absent from inception. The dicliny is established at different development stages: stamens or carpels are absent from inception (Cannabis sativa), initiated and aborted during early (Trema micrantha, before sporo/gametogenesis) or late (Celtis iguanaea, after sporo/gametogenesis) development. Furthermore, in all species studied the carpels are congenitally united and the pseudomonomerous nature of the gynoecium is confirmed. Glandular trichomes are distributed on the bracts, sepals, anther connective and receptacle. Special floral features shared by species of Cannabaceae include precocious ovule development and sepals that are each vascularized by one bundle. The reduced flowers of Cannabaceae are the result of the absence from inception and/or abortion of organs and even of a whole whorl at different developmental stages, which were probably selected in response to pressures exerted by the similar pollination mechanism.

It is not a disaster: molecular and morphologically based phylogenetic analysis of Rondeletieae and the Rondeletia complex (Cinchonoideae, Rubiaceae)

The circumscription of Rondeletieae has changed considerably over time. Historically, 85 genera have been included, characterized by small trees, imbricate corolla aestivation and capsules. Molecular studies led to a narrower circumscription and stressed out that Rondeletieae is mainly an Antillean clade. As historically circumscribed, Rondeletieae is both polyphyletic (involving several tribes, including Condamineae, Sipaneeae, etc.) and paraphyletic with respect to Guettardeae. Uncertainties persist on the tribal circumscription, and its generic limits and relationships. We report results of phylogenetic analyses under parsimony, maximum likelihood and Bayesian analyses including 179 taxa, using concatenated datasets of three chloroplast DNA regions (petD, trnL–F and trnK–matK), their DNA microstructural characters as well as morphology. Our objectives are to: test the monophyly of Rondeletieae and evaluate phylogenetic relationships, evaluate potential characters for tribal and generic delimitation and propose a circumscription for Rondeletieae according to phylogenetic hypothesis using morphological characters. The tribes Guettardeae and Rondeletieae are strongly supported as sister: quincuncial aestivation and spathulate corolla lobes are proposed as a unique combination of characters supporting this relationship. We did not find synapomorphies for each tribe, but there is a set of character combinations that allows distinguishing both tribes that includes the corolla and pollen ornamentation as well as fruit and seed morphology. After evaluating other genera not included in the phylogenies, our proposal is that Rondeletieae contains approximately 120 species in 20 neotropical genera: Acrosynanthus, Acunaeanthus, Blepharidium, Donnellyanthus, Mazaea, Phyllomelia, Rachicallis, Roigella, Rondeletia, Rovaeanthus, Suberanthus, Tainus, plus one tentatively included and recently described (Jamaicanthus).

What is the nature of petals in Caryophyllaceae? Developmental evidence clarifies their evolutionary origin.

Floral development is a powerful tool to infer homologies of floral organs and to understand floral evolution. Caryophyllaceae is a major family of core Caryophyllales that possesses petal-like structures (petaloids) with a great diversity in shape. The main purpose of this study is to determine the nature of the second whorl of floral organs in Caryophyllaceae. Mainstream views consider ancestors of Caryophyllaceae as apetalous and interpret petals as centrifugally derived staminodial appendages. This hypothesis, based on morphological similarities of petals with stamens and previous ancestral state reconstruction, is tested here. A floral developmental investigation of five species was carried out using scanning electron microscopy, combined with character optimization of the presence or absence of second-whorl petaloids. The calyx is always well developed with a quincuncial aestivation. Petaloids either develop by fractionation of common stamen-petal primordia, as in Spergularia, or petaloid development is independent and precedes alternisepalous stamens in Saponaria and Sagina. In Sagina the petaloid whorl is always fully formed but alternisepalous stamens are often reduced or missing. Petaloids are absent in Gymnocarpos and the investigated Cerastium. Developmental evidence and character mapping reject the hypothesis that petaloids represent a staminodial whorl and suggest that they are independent structures equivalent to second-whorl petals of most Pentapetalae and present in the basal Caryophyllaceae. Heterochronic shifts, including a delay in petal development and acceleration of androecial growth, are responsible for the amalgamation of petals with the androecium as common stamen-petal primordia and their appearance as stamen-derived appendages. Selective pollinator pressure in Caryophyllaceae led to variable petal expansion or reduction and loss. This trend corresponds largely with the general tendency in the core Caryophyllales for petal loss and perianth reorganization.

Molecular phylogeny of Koenigia L. (Polygonaceae: Persicarieae): Implications for classification, character evolution and biogeography

To examine the phylogenetic relationships of Koenigia sensu lato (Polygonaceae), 43 samples representing all species of Koenigia and closely related taxa (e.g., Aconogonon, Bistorta, and Persicaria) were sequenced for nuclear ITS and four plastid regions (trnL-F, atpB-rbcL, rbcL, and rpl32-trnL(UAG)). Phylogenetic analyses indicate that Koenigia recognized by Hedberg is paraphyletic and that the basal species K. delicatula should be reassigned to a separate new genus. Based on these findings, we further propose that the genus Koenigia sensu lato be circumscribed to include five species. Ancestral state reconstruction showed that the pollen apertures likely evolved in parallel in the Aconogonon–Koenigia–Bistorta clade and Persicaria clade and that tricolpate pollen is most likely to be the ancestral state. Quincuncial aestivation likely evolved during the early evolution of Koenigia and its close relatives. Our findings suggest that the uplift of the Himalayas has played an important role in promoting species diversification of Koenigia. Koenigia islandica expanded its range during Pleistocene glacial cycles by tracking changes in newly available habitats.

A new species of <i>Croton</i> section <i>Cleodora</i> (Euphorbiaceae s.s.) from Minas Gerais, Brazil

Croton stellatoferrugineus, a new species from Brazil, is here described and illustrated. This new species is endemic to dry forests at the base of “Pico do Itambé”, the highest point of the Espinhaço range, in Minas Gerais State, occurring at 700–900 m elevation. Croton stellatoferrugineus presents morphological features consistent with its inclusion in Croton section Cleodora, due to its shrubby habit, petiolar basilaminar glands, 15 stamens, pistillate flowers with quincuncial aestivation, and multifid styles connate at the base.

Glandulocalyx upatoiensis, a fossil flower of Ericales (Actinidiaceae/Clethraceae) from the Late Cretaceous (Santonian) of Georgia, USA.

Ericales are a major group of extant asterid angiosperms that are well represented in the Late Cretaceous fossil record, mainly by flowers, fruits and seeds. Exceptionally well preserved fossil flowers, here described as Glandulocalyx upatoiensis gen. & sp. nov., from the Santonian of Georgia, USA, yield new detailed evidence of floral structure in one of these early members of Ericales and provide a secure basis for comparison with extant taxa. The floral structure of several fossil specimens was studied by scanning electron microscopy (SEM), light microscopy of microtome thin sections and synchrotron-radiation X-ray tomographic microscopy (SRXTM). For direct comparisons with flowers of extant Ericales, selected floral features of Actinidiaceae and Clethraceae were studied with SEM. Flowers of G. upatoiensis have five sepals with quincuncial aestivation, five free petals with quincuncial aestivation, 20-28 stamens arranged in a single series, extrorse anther orientation in the bud, ventral anther attachment and a tricarpellate, syncarpous ovary with three free styles and numerous small ovules on axile, protruding-diffuse and pendant placentae. The calyx is characterized by a conspicuous indumentum of large, densely arranged, multicellular and possibly glandular trichomes. Comparison with extant taxa provides clear evidence for a relationship with core Ericales comprised of the extant families Actinidiaceae, Roridulaceae, Sarraceniaceae, Clethraceae, Cyrillaceae and Ericaceae. Within this group, the most marked similarities are with extant Actinidiaceae and, to a lesser degree, with Clethraceae. More detailed analyses of the relationships of Glandulocalyx and other Ericales from the Late Cretaceous will require an improved understanding of the morphological features that diagnose particular extant groups defined on the basis of molecular data.

Hekkingia (Violaceae), a New Arborescent Violet Genus from French Guiana, with a Key to Genera in the Family

Abstract A new genus of arborescent violet, Hekkingia, is described from Neotropical lowland rainforest near Cayenne in northeastern French Guiana. Racemose inflorescences, fully fused filaments and rudimentary dorsal connective appendages place the only known species, Hekkingia bordenavei, near the genus Paypayrola, while strongly tuberculate capsules ally it also to Amphirrhox, the two genera comprising the Subtribe Paypayrolinae to which the new genus is tentatively assigned. Hekkingia shares scarcely differentiated corollas with quincuncial aestivation, very large obovate sepals, red to pink capsules and carunculate seeds with few other genera (but not in this combination), whereas its basally cauliflorous habit, very large bracts and pedicel bractlets are unique in the family. A key to genera of the Violaceae provides a context for this French Guianan novelty.

RbcL Phylogeny and Character Evolution in Polygonaceae

Abstract Phylogenetic relationships within Polygonaceae, with an emphasis on Polygonum and segregate genera (e.g., Emex, Persicaria, Polygonella), are investigated using sequence data from the chloroplast gene rbcL. The results show the currently recognized subfamilies are likely not monophyletic. Polygonum as currently recognized is paraphyletic. Three strongly supported clades of Polygonum s.l. were found. Contained within these three clades of Polygonum s.l. are representatives of Atraphaxis, Bistorta, Koenigia and Persicaria. Other strongly supported clades found are Coccoloba, Eriogonum, Fagopyrum, Rumex and Triplaris. Patterns of character evolution were studied by mapping characters onto the phylogeny. A woody habit likely evolved multiple times within the family (in Atraphaxis, Koenigia, Muehlenbeckia, Polygonella and in the ancestor to the Eriogonum–Brunnichia clade). Quincuncial aestivation likely evolved early in the history of the family with four subsequent losses in the Rumex–Oxyria clade, M...