Floral morphology and development reveal extreme diversification in some species of Croton (Euphorbiaceae).

Floral diversity of Croton, the second largest genus in Euphorbiaceae, is currently under-explored. Several clades demonstrate an unusual floral morphology, e.g., lower or higher stamen number, bilateral symmetry and reduced ovary, but have never been investigated in a comparative study with typical Croton. This study examined morphology and ontogeny of flowers in nine Croton species from different clades within the genus with light and scanning microscopy, resin sectioning and micro-computed tomography. In staminate flowers, great variations of stamen number and arrangement are observed. The ancestral androecium likely consisted of two or more whorls with the outermost antepetalous stamen whorl developing centrifugally. Modification by reduction of the antepetalous whorl resulted in an outer alternipetalous stamen whorl in Croton section Moacroton, subgenus Quadrilobi. Several species in the subgenus Geiseleria show an independent reduction of stamen numbers by absence of a centrifugal development with the antepetalous whorl the first whorl to develop. Petal losses are observed in the distantly related C. setiger and C. dioicus. Chaotic stamen arrangement is found in C. celtidifolius (subgenus Adenophylli) as a result of a secondary stamen increase. In pistillate flowers, reduction of carpel numbers happened three times in the subgenus Geiseleria. C. monanthogynus has a bicarpellate ovary, while in C. setiger and C. michauxii the ovary is monocarpellate. Reduction of carpel number is linked with merism change and perianth reduction. The ovary in C. michauxii has basal placentation which is unique among all Croton. Moreover, strong bilateral sepals and nectaries are observed in species from section Julocroton. Therefore, the floral diversity of some species in the genus Croton could be explained by developmental modification of an ancestral form via reduction, rearrangement of stamen whorls, and symmetry shifts.

Leaf plasticity across wet and dry seasons in Croton blanchetianus (Euphorbiaceae) at a tropical dry forest.

Plant species of the Brazilian Caatinga experience seasonal wet and dry extremes, requiring seasonally different leaf characteristics for optimizing water availability. We investigated if Croton blanchetianus Baill exhibits leaf morphoanatomical traits across seasons and positioning in sunlight/natural shade. Leaves of ten 1-3 m tall plants in full sunlight and ten in natural shade were assessed in May, July (wet season), October and December (dry season) 2015 for gas exchange, leaf size, lamina and midrib cross sections (14 parameters), and chloroplast structure (5 parameters). Net photosynthesis was greater during the wet season (21.6 µm-2 s-1) compared to the dry season (5.8 µm-2 s-1) and was strongly correlated with almost all measured parameters (p < 0.01). Shaded leaves in the wet season had higher specific leaf area (19.9 m2 kg-1 in full-sun and 23.1 m2 kg-1 in shade), but in the dry season they did not differ from those in full sun (7.5 m2 kg-1 and 7.2 m2 kg-1). In the wet season, the expansion of the adaxial epidermis and mesophyll lead to larger and thicker photosynthetic area of leaves. Furthermore, chloroplast thickness, length and area were also significantly larger in full sunlight (2.1 µm, 5.1 µm, 15.2 µm2; respectively) and shaded plants (2.0 µm, 5.2 µm, 14.8 µm2; respectively) during wetter months. Croton blanchetianus exhibits seasonal plasticity in leaf structure, presumably to optimize water use efficiency during seasons of water abundance and deficit. These results suggest that the species is adaptable to the increased drought stress projected by climate change scenarios.

Force of habit: shrubs, trees and contingent evolution of wood anatomical diversity using Croton (Euphorbiaceae) as a model system.

Background and Aims: Wood is a major innovation of land plants, and is usually a central component of the body plan for two major plant habits: shrubs and trees. Wood anatomical syndromes vary between shrubs and trees, but no prior work has explicitly evaluated the contingent evolution of wood anatomical diversity in the context of these plant habits. Methods: Phylogenetic comparative methods were used to test for contingent evolution of habit, habitat and wood anatomy in the mega-diverse genus Croton (Euphorbiaceae), across the largest and most complete molecular phylogeny of the genus to date. Key Results: Plant habit and habitat are highly correlated, but most wood anatomical features correlate more strongly with habit. The ancestral Croton was reconstructed as a tree, the wood of which is inferred to have absent or indistinct growth rings, confluent-like axial parenchyma, procumbent ray cells and disjunctive ray parenchyma cell walls. The taxa sampled showed multiple independent origins of the shrub habit in Croton , and this habit shift is contingent on several wood anatomical features (e.g. similar vessel-ray pits, thick fibre walls, perforated ray cells). The only wood anatomical trait correlated with habitat and not habit was the presence of helical thickenings in the vessel elements of mesic Croton . Conclusions: Plant functional traits, individually or in suites, are responses to multiple and often confounding contexts in evolution. By establishing an explicit contingent evolutionary framework, the interplay between habit, habitat and wood anatomical diversity was dissected in the genus Croton . Both habit and habitat influence the evolution of wood anatomical characters, and conversely, the wood anatomy of lineages can affect shifts in plant habit and habitat. This study hypothesizes novel putatively functional trait associations in woody plant structure that could be further tested in a variety of other taxa.

Germination of Croton urucurana L. seeds exposed to different storage temperatures and pre-germinative treatments.

The present work evaluated the germinability and vigor of Croton urucurana seeds. 1) Seeds were sorted by color (caramel, gray and black) and were subjected to seven different pre-germination treatments followed by incubation at 20ºC, 25°C or 20/30°C. 2) Seeds were stored in cold chambers or at room temperature for up to 300 days and were subsequently incubated at 20/30ºC in a germination chamber or under greenhouse conditions. Only gray seeds showed significant germination rates. The highest first count percentages of total germination and the highest germination speed indices were observed in control seeds and in those which were treated with water or 200 mg.L(-1) gibberellic acid for 12 hours. Seeds stored under refrigeration showed the highest values for all of the characteristics examined, as well as less electrical conductivity of the imbibing solution. Seedlings were more vigorous when seeds were stored for 300 days in a cold chamber. The seedlings production can be increased by incubating the seeds at alternating temperatures (20/30°C). The seeds do not need pre-germination treatments.

Molecular phylogenetics and character evolution of the "sacaca" clade: novel relationships of Croton section Cleodora (Euphorbiaceae).

Phylogenetic relationships of Croton section Cleodora (Klotzsch) Baill. were evaluated using the nuclear ribosomal ITS and the chloroplast trnL-F and trnH-psbA regions. Our results show a strongly supported clade containing most previously recognized section Cleodora species, plus some other species morphologically similar to them. Two morphological synapomorphies that support section Cleodora as a clade include pistillate flowers in which the sepals overlap to some degree, and styles that are connate at the base to varying degrees. The evolution of vegetative and floral characters that have previously been relied on for taxonomic decisions within this group are evaluated in light of the phylogenetic hypotheses. Within section Cleodora there are two well-supported clades, which are proposed here as subsections (subsection Sphaerogyni and subsection Spruceani). The resulting phylogenetic hypothesis identifies the closest relatives of the medicinally important and essential oil-rich Croton cajucara Benth. as candidates for future screening in phytochemical and pharmacological studies.

Anodic asymmetry of leaves and flowers and its relationship to phyllotaxis.

BACKGROUND AND AIMS: New approaches are needed to evaluate the various hypotheses of phyllotaxis, and an examination of anodic leaf asymmetry may be one such approach. METHODS: Data were collected on the direction of midrib curvature and leaf coil in Syngonium podophyllum, the location of floral buds in Acalypha virginica, the position of secondary leaves of Croton variegatus 'Banana' and the relative size of half-lamina in Aglaonema crispum and Calathea ornata. KEY RESULTS: All five features were exclusively anodic with respect to the direction of the genetic spiral regardless of whether the spiral was clockwise or counterclockwise. CONCLUSIONS: Any phyllotactic mechanism must include some asymmetric component which cannot be explained by the prevalent hypotheses of contact parastichies, inhibitory fields, available space, pressure waves and auxin transport. The most favourable hypothesis is the primary vasculature explanation as it includes an asymmetric feature.