Is Santaniella a ranunculid? Reassessment of this enigmatic fossil angiosperm from the Lower Cretaceous (Aptian, Crato Konservat-Lagerstätte, Brazil) provides a new interpretation.

PREMISE: The Lower Cretaceous Crato Konservat-Lagerstätte (CKL) preserves a rich flora that includes early angiosperms from northern Gondwana. From this area, the recently described fossil genus Santaniella was interpreted as a ranunculid (presumably Ranunculaceae). However, based on our examination of an additional specimen and a new phylogenetic analysis, we offer an alternative interpretation. METHODS: The new fossil was collected from an active quarry for paving stones in the state of Ceará, northeastern Brazil. We assessed support for alternative phylogenetic hypotheses using a combined analysis of morphological data and DNA sequence data using Bayesian inference. We used a consensus network to visualize the posterior distribution of trees, and we used RoguePlot to illustrate the support for alternative positions on a scaffold tree. RESULTS: The new material includes a flower-like structure not present in the original material and also includes follicles preserved at early stages of development. The flower-like structure is a compact terminal cluster of elliptical sterile laminar organs surrounding internal filamentous structures that occur on flexuous axes. Phylogenetic analyses did not support the fossil placement among eudicots. Instead, Santaniella appears to belong in the magnoliid clade. CONCLUSIONS: The presence of seeds in a marginal-linear placentation and enclosed in a follicle supports the fossil as an angiosperm. However, even though most characters are clearly recognizable, its combination of characters does not provide strong support for a close relationship to any extant order of flowering plants. Its position in the magnoliid clade is intriguing and, based on plicate carpels, it is definitely a mesangiosperm.

Floral development and vasculature in Eriocaulon (Eriocaulaceae) provide insights into the evolution of Poales.

BACKGROUND AND AIMS: Floral developmental studies are crucial for understanding the evolution of floral structures and sexual systems in angiosperms. Within the monocot order Poales, both subfamilies of Eriocaulaceae have unisexual flowers bearing unusual nectaries. Few previous studies have investigated floral development in subfamily Eriocauloideae, which includes the large, diverse and widespread genus Eriocaulon. To understand floral variation and the evolution of the androecium, gynoecium and floral nectaries of Eriocaulaceae, we analysed floral development and vasculature in Eriocaulon and compared it with that of subfamily Paepalanthoideae and the related family Xyridaceae in a phylogenetic context. METHODS: Thirteen species of Eriocaulon were studied. Developmental analysis was carried out using scanning electron microscopy, and vasculature analysis was carried out using light microscopy. Fresh material was also analysed using scanning electron microscopy with a cryo function. Character evolution was reconstructed over well-resolved phylogenies. KEY RESULTS: Perianth reductions can occur due to delayed development that can also result in loss of the vascular bundles of the median sepals. Nectariferous petal glands cease development and remain vestigial in some species. In staminate flowers, the inner stamens can emerge before the outer ones, and carpels are transformed into nectariferous carpellodes. In pistillate flowers, stamens are reduced to staminodes and the gynoecium has dorsal stigmas. CONCLUSIONS: Floral morphology is highly diverse in Eriocaulon, as a result of fusion, reduction or loss of perianth parts. The nectariferous carpellodes of staminate flowers originated first in the ancestor of Eriocaulaceae; petal glands and nectariferous branches of pistillate flowers originated independently in Eriocaulaceae through transfer of function. We present a hypothesis of floral evolution for the family, illustrating a shift from bisexuality to unisexuality and the evolution of nectaries in a complex monocot family, which can contribute to future studies on reproductive biology and floral evolution in other groups.

Development of the permanent tetrad wall in Juncus L. (Juncaceae, Poales).

Juncaceae, a cosmopolitan family, belong to the cyperid clade of Poales together with Cyperaceae and Thurniaceae. Pollen grain of Juncaceae, as in Thurniaceae, is dispersed in a permanent tetrad, and knowledge about the ontogeny of its wall is still incipient, based on data from only one species. This study aims to analyze the formation of the pollen wall of seven Juncus species in order to characterize the timing and the ontogenetic events that lead to the cohesion of the four pollen grains in a permanent tetrad. Anthers at different developmental stages were submitted to techniques of light microscopy and transmission electron microscopy; dehiscent anthers with mature pollens were also analyzed in scanning electron microscopy. In all the species here studied, callose deposits around each microsporocyte, with dissolution prior to meiosis. Microspore wall starts at the end of the second meiotic division with formation of primexine. Exine comprises tectum, columellae, and foot layer. During cytokinesis, cell plates form the internal wall of the pollen tetrad. Mature permanent tetrad is enveloped externally by both the exine and intine and internally by the intine and the foot layer, which forms the continuous internal wall. Callose was detected in the early stages of microsporocytes, although reported to be absent in Juncaceae. Our data confirm the variation in Juncaceae cytokinesis and the occurrence of simple cohesion due to the presence of a continuous tectum along the pollen tetrad.

Release of developmental constraints on tetrad shape is confirmed in inaperturate pollen of Potamogeton.

BACKGROUND AND AIMS: Microsporogenesis in monocots is often characterized by successive cytokinesis with centrifugal cell plate formation. Pollen grains in monocots are predominantly monosulcate, but variation occurs, including the lack of apertures. The aperture pattern can be determined by microsporogenesis features such as the tetrad shape and the last sites of callose deposition among the microspores. Potamogeton belongs to the early divergent Potamogetonaceae and possesses inaperturate pollen, a type of pollen for which it has been suggested that there is a release of the constraint on tetrad shape. This study aimed to investigate the microsporogenesis and the ultrastructure of pollen wall in species of Potamogeton in order to better understand the relationship between microsporogenesis features and the inaperturate condition. METHODS: The microsporogenesis was investigated using both light and epifluorescence microscopy. The ultrastructure of the pollen grain was studied using transmission electron microscopy. KEY RESULTS: The cytokinesis is successive and formation of the intersporal callose wall is achieved by centrifugal cell plates, as a one-step process. The microspore tetrads were tetragonal, decussate, T-shaped and linear, except in P. pusillus, which showed less variation. This species also showed a callose ring in the microsporocyte, and some rhomboidal tetrads. In the mature pollen, the thickening observed in a broad area of the intine was here interpreted as an artefact. CONCLUSIONS: The data support the view that there is a correlation between the inaperturate pollen production and the release of constraint on tetrad shape. However, in P. pusillus the tetrad shape may be constrained by a callose ring. It is also suggested that the lack of apertures in the pollen of Potamogeton may be due to the lack of specific sites on which callose deposition is completed. Moreover, inaperturate pollen of Potamogeton would be better classified as omniaperturate.

Floral development and vascularization help to explain merism evolution in Paepalanthus (Eriocaulaceae, Poales).

BACKGROUND: Flowers in Eriocaulaceae, a monocot family that is highly diversified in Brazil, are generally trimerous, but dimerous flowers occur in Paepalanthus and a few other genera. The floral merism in an evolutionary context, however, is unclear. Paepalanthus encompasses significant morphological variation leading to a still unresolved infrageneric classification. Ontogenetic comparative studies of infrageneric groups in Paepalanthus and in Eriocaulaceae are lacking, albeit necessary to establish evolution of characters such as floral merism and their role as putative synapomorphies. METHODS: We studied the floral development and vascularization of eight species of Paepalanthus that belong to distinct clades in which dimery occurs, using light and scanning electron microscopies. RESULTS: Floral ontogeny in dimerous Paepalanthus shows lateral sepals emerging simultaneously and late-developing petals. The outer whorl of stamens is absent in all flowers examined here. The inner whorl of stamens becomes functional in staminate flowers and is reduced to staminodes in the pistillate ones. In pistillate flowers, vascular bundles reach the staminodes. Ovary vascularization shows ventral bundles in a commissural position reaching the synascidiate portion of the carpels. Three gynoecial patterns are described for the studied species: (1) gynoecium with a short style, two nectariferous branches and two long stigmatic branches, in most species; (2) gynoecium with a long style, two nectariferous branches and two short stigmatic branches, in P. echinoides; and (3) gynoecium with long style, absent nectariferous branches and two short stigmatic branches, in P. scleranthus. DISCUSSION: Floral development of the studied species corroborates the hypothesis that the sepals of dimerous flowers of Paepalanthus correspond to the lateral sepals of trimerous flowers. The position and vascularization of floral parts also show that, during dimery evolution in Paepalanthus, a flower sector comprising the adaxial median sepal, a lateral petal, a lateral stamen and the adaxial median carpel was lost. In the staminate flower, the outer whorl of staminodes, previously reported by different authors, is correctly described as the apical portion of the petals and the pistillodes are reinterpreted as carpellodes. The occurrence of fused stigmatic branches and protected nectariferous carpellodes substantiates a close relationship between P. sect. Conodiscus and P. subg. Thelxinoë. Free stigmatic branches and exposed carpellodes substantiate a close relationship between P. sect. Diphyomene, P. sect. Eriocaulopsis and P. ser. Dimeri. Furthermore, the loss of nectariferous branches may have occurred later than the fusion of stigmatic branches in the clade that groups P. subg. Thelxinoë and P. sect. Conodiscus.

Importance of anatomical leaf features for characterization of three species of Mapania (Mapanioideae, Cyperaceae) from the Amazon Forest, Brazil / Importância dos caracteres anatômicos foliares na caracterização de três espécies de Mapania (Mapanioideae, Cyperaceae) da Floresta Amazônica, Brasil

Mapania belongs to Mapanioideae, a quite controversial subfamily in Cyperaceae due to the existence of unusual characters in both reproductive and vegetative organs. The genus is represented by seven species in Northern Brazil but taxonomic valuable information related to the leaf organs is still unknown. The present study aimed the anatomical description of the leaf organs (either basal leaves or cataphylls and involucral bracts) of three representative Brazilian species of Mapania. Samples of cataphylls, basal leaves and involucral bracts were sectioned and stained for observations under light microscopy. The involucral bracts provide the most elucidative characters (ten) to distinguish the three species The basal leaves provides six distinguishing characters and are useful to M. macrophylla and M. pycnostachya, as they are absent in M. sylvatica. Mesophyll arrangement in the involucral bracts supports the circumscription of M. macrophylla and M. pycnostachya in M. sect. Pycnocephala and of M. sylvatica in M. sect. Mapania. Some features as thin-walled epidermal cells, stomata level and aerenchyma were considered to be adaptive to the humid environment in which the species occur. The translucent cells are here considered as aerenchyma precursors and a supportive function is assumed for the bulliform cells on the basal leaves and involucral bracts. No silica bodies were found which confirm it as a diagnostic character of Mapania among Hypolytreae genera.

Mapania pertence à subfamília Mapanioideae que apresenta caracteres incomuns às demais Cyperaceae, tanto vegetativos como reprodutivos. O gênero é representado por sete espécies no norte do Brasil e apresenta ainda lacunas de informações especialmente relacionadas aos órgãos foliares. O presente estudo objetivou a descrição anatômica dos órgãos foliares (folhas basais, ou catafilos, e brácteas involucrais) de três espécies representativas de Mapania no território brasileiro. Amostras de catafilos, folhas basais e brácteas involucrais foram seccionadas e coradas para observações em microscopia de luz. As brácteas involucrais foram as mais elucidativas na separação das espécies, com dez caracteres variáveis, uma vez que ocorrem em todas as espécies durante o estágio reprodutivo. As folhas basais forneceram seis caracteres úteis na distinção de M. macrophylla e M. pycnostachya, já que não ocorrem em M. sylvatica. seguidas das folhas basais. O arranjo do mesofilo das brácteas involucrais suporta a circunscrição de M. macrophylla e M. pycnostachya em M. sect. Pycnocephala e de M. sylvatica em M. sect. Mapania. Características adaptativas ao ambiente úmido também foram encontradas, como epiderme com paredes delgadas, níveis dos estômatos e presença de aerênquima. As células translúcidas do mesofilo foram descritas como precursoras de aerênquima e considerou-se que as células buliformes presentes nas folhas e brácteas involucrais apresentam função de suporte desses órgãos. Corpos silicosos não foram encontrados sustentando o seu uso na delimitação de Mapania e dos demais gêneros de Hypolytreae.

Leaf Morphological Plasticity of Tree Species from Two Developmental Stages in Araucaria Forest

This study compared the morphological and anatomical variations of the leaves of four shade-tolerant tree species Allophylus edulis (St.-Hil.) Radlk (Sapindaceae), Casearia sylvestris Sw. (Salicaceae), Cupania vernalis Cambess. (Sapindaceae) and Luehea divaricata Mart. (Malvaceae) from a fragment of Araucaria forest in two developmental stages. Morphological and anatomical traits, such as leaf and tissue thickness, leaf area, leaf dry mass, specific leaf area, leaf density and stomata density were measured from 30 leaves of each developmental stage. The phenotypic plasticity index was also calculated for each quantitative trait. The results showed that the four species presented higher mean values ​​for specific leaf area and spongy/palisade parenchyma ratio at young stage, and higher mean values ​​for stomata density, total and palisade parenchyma thickness in the adult stage. The plasticity index demonstrated that L. divricata presented highest plasticity for both the morphological and anatomical traits while A. edulis displayed the lowest plasticity index. The results of this study indicated that the leaves of these species exhibited distinct morphological traits at each stage of development to cope with acting environmental factors.

Seed morphology and post-seminal development of Tillandsia L. (Bromeliaceae) from the "Campos Gerais", Paraná, Southern Brazil

This work studied the morphology and post-seminal development of seeds of Tillandsia gardneri Lindl., T. streptocarpa Baker, and T. stricta Sol. ex Sims (Bromeliaceae) from the "Campos Gerais", State of Paraná, Southern Brazil. The seeds (ca. 2.0 mm long) presented plumose appendages at their micropylar end, an important structure in the taxonomy of the family since it represented an elaborate apparatus allowing effective wind-dispersal. Seeds germinated after circa twenty days. The haustorial cotyledon, responsible for the seedling nutrition, appeared first during the germination process. No cataphyll or primary roots developed. A small tank came along in the seedling, formed by the base of the sheath of the first leaf, which overlapped that of the second leaf. This tank can store water and litter. The adventitious roots developed later.

Foi estudada a morfologia da semente e do desenvolvimento pós-seminal de Tillandsia gardneri Lindl., T. streptocarpa Baker, e T. stricta Sol. ex Sims (Bromeliaceae), que ocorrem nos Campos Gerais do Paraná, Brasil. As sementes apresentaram cerca de 2,0 mm de comprimento e testa com apêndices plumosos na região micropylar. A presença desses apêndices é um caráter importante na taxonomia da família e garante sucesso na dispersão anemocórica das sementes. Após a embebição, as sementes levaram cerca de 20 dias para germinar. A primeira estrutura que emergiu durante a germinação foi o cotilédone haustorial, que é responsável pela nutrição da plântula. Eram ausentes catafilo e raiz primária. Na plântula ocorreu a formação de um pequeno tanque, que é constituído pela base da bainha da folha primária superposta à base da bainha da folha secundária. Esse pequeno tanque permaneceu na posição vertical e armazenou água e detritos orgânicos para a nutrição da plântula. As raízes adventícias emergiram tardiamente.