Patterns of sex expression and sex ratios are key features of the life histories of organisms. Bryophytes are the only haploid-dominant land plants. In contrast with seed plants, more than half of bryophyte species are dioecious, with rare sexual expression and sporophyte formation and a commonly female-biased sex ratio. We asked whether variation in sex expression, sex ratio and sporophyte frequency in ten dioecious pleurocarpous wetland mosses of two different families was best explained by assuming that character states evolved: (1) in ancestors within the respective families or (2) at the species level as a response to recent habitat conditions. Lasso regression shrinkage identified relationships between family membership and sex ratio and sporophyte frequency, whereas environmental conditions were not correlated with any investigated reproductive trait. Sex ratio and sporophyte frequency were correlated with each other. Our results suggest that ancestry is more important than the current environment in explaining reproductive patterns at and above the species level in the studied wetland mosses, and that mechanisms controlling sex ratio and sporophyte frequency are phylogenetically conserved. Obviously, ancestry should be considered in the study of reproductive character state variation in plants.
Sex ratio variation is common among organisms with separate sexes. In bryophytes, sex chromosome segregation at meiosis suggests a balanced progeny sex ratio. However, most bryophyte populations exhibit female-biased phenotypic sex ratios based on the presence of reproductive structures on gametophytes. Many bryophyte populations do not form sexual organs, and genotypic sex ratio variation in such populations is mostly unknown. We tested sex expression, and phenotypic and genotypic sex ratios against environmental parameters in natural populations of the unisexual wetland moss Drepanocladus lycopodiodes at 11 sites in each of three regions in southern Sweden. We identified sex in 660 individual ramets, based on sexual structures, when present, or with a specifically designed molecular marker, when absent. All regions exhibited a female bias in phenotypic and genotypic sex atios. Sex ratio biases and sex expression differed between regions. Sex ratios were less male-biased in larger patches. Wetter patches exhibited a stronger female bias in genotypic sex ratio and lower sex expression. This is the first evidence of environmental effects on genotypic sex ratio in mosses. A higher frequency of females in wet patches could be due to higher female resource demands for sporophyte production or higher male sensitivity to wetness. A higher incidence of females than males in moister sites aligns with female flowering plants, but differs from reproductive bryophytes in drier environments. Taken together with previous results, our data indicate that sex ratio variation and its drivers differ among species, their life histories and environments.
Anatolia was a crossroads for mammal migration during the Miocene due to intermittent land connections between Africa and Anatolia and persisting warm conditions. Here, we investigated a palynological section from middle Miocene sediments of Eskihisar (southwestern Anatolia) in order to establish biogeographic links of the palynoflora and to infer the palaeoenvironment. Four algal palynomorphs, nine spore taxa, eight gymnosperms, three monocots, and 67 dicot pollen types were encountered and investigated using the “single grain method” that combines light microscopy and scanning electron microscopy. Two pollen zones reflect different phases of basin development. Zonal vegetation remained fairly stable across the section and reflects heterogeneous environments including broad-leaved deciduous forest, subtropical forest, and sclerophyllous and semi-evergreen oak forest. Conifers were accessory elements in the broad-leaved deciduous forest communities and replaced these at higher elevations. Some herbaceous taxa (Plumbaginaceae) indicate scattered occurrences of sandy and/or rocky soils. Biogeographic affinities are general Northern Hemispheric, North American, and East Asian as also suggested by the macro fossil record. Only two taxa provide potential biogeographic links with the African flora. This suggests that biome shifts of plant taxa between African subtropical /tropical biomes and Anatolian (western Eurasian) temperate forests and shrublands may have been rare in the middle Miocene.
Generic limits of Chrysophyllum and Pouteria (Chrysophylloideae, Sapotaceae) have been found to be untenable. We here search for natural lineages in Neotropical Chrysophylloideae by sampling 101 terminals for molecular sequences of nuclear ribosomal DNA (external and internal transcribed spacer), the nuclear gene RPB2 and 17 morphological characters. Data were analysed with Bayesian inference and parsimony jackknifing. Morphological traits were finally optimized onto the tree to identify the most coherent characters. The resulting phylogenetic tree suggests that the limits of the well-known genera Chrysophyllum and Pouteria must be amended. Diploon, Ecclinusa and Elaeoluma can be maintained and Chrysophyllum sections Ragala section Prieurella and the satellite gen- era Achrouteria, Cornuella, Martiusella and Nemaluma merit generic resurrection. Lucuma may be restored if the type species belongs to the clade. The accepted genera Chromolucuma, Pradosia and Sarcaulus gain strong clade support, but are embedded in a core clade of Pouteria and may be relegated to the subgeneric level if morphologi- cal studies cannot provide evidence concurring with narrow generic concepts. Circumscriptions of Micropholis and Chrysophyllum sections Chrysophyllum and Villocuspis remain unclear and must be explored by using an extended taxon sampling. We predict that yet-to-be-analysed species of Pouteria sections Franchetella, Gayella, Oxythece and Pouteria and members of the currently accepted genera Chromolucuma, Pradosia and Sarcaulus will fall inside the core clade of Pouteria when analysed.
Sapotaceae belongs to the heterogeneous order Ericales and exhibits extensive diversity in floral morphology. Although pentamery is widespread and probably the ancestral condition, some clades are extremely variable in merism, with fluctuations between tetramery to hexamery and octomery, affecting different floral organs to different degrees. We assessed the different states of merism in Sapotaceae to determine the evolution of this character among different clades. The floral morphology and development of nine species from eight genera were investigated using scanning electron microscopy (SEM). Furthermore, floral characters related to merism were mapped onto a phylogenetic tree to analyse the distribution and evolutionary significance of merism in the family. Developmental evidence shows that changes in merism are linked to a concerted multiplication of organs among whorls and an increase in whorls through the displacement of organs. Although pentamery is reconstructed as the ancestral condition, a reduction to tetramery or an increase to a higher merism (mainly hexamery or octomery) has evolved at least five times in the family. Fluctuations in merism between different whorls are not random but occur in a coordinated pattern, presenting strong synapomorphies for selected clades. Octomery has evolved at least twice, in Isonandreae from tetramery and in Sapoteae-Mimusopinae from pentamery. Hexamery has evolved at least three times, independently in Northia, the Palaquium clade of Isonandreae and derived from octomery in Sapoteae-Mimusopinae. Three possibilities of merism increase have been identified in Sapotaceae: (1) a concerted increase affecting all organs more or less equally (Palaquium clade of Isonandreae, Sapoteae); (2) a coordinated increase in petals, stamens and mostly carpels without effect on sepals (Labourdonnaisia, Payena–Madhuca clade of Isonandreae); (3) an increase in carpels independently of other organs (Burckella, Letestua, Labramia, etc.). A major shift affecting all Sapotaceae, except Isonandreae, is the sterilization or loss of the antesepalous stamen whorl. The presence of two fertile stamen whorls in Isonandreae indicates a possible reversal or a retained plesiomorphy. In a number of genera, stamens are secondarily increased independently of changes in merism. Descriptions of flowers listing only organ numbers are thus misleading in the inference of evolutionary relationships, as they do not differentiate between changes in merism affecting the number of perianth whorls and other changes affecting the androecium, such as sterilization, loss or occasional doubling of antepetalous stamens.
Faramea is characterized by white or blue, tetramerous corollas and blue-black, fleshy fruits with a single, large pyrene. Both infrageneric relationships and species boundaries are poorly understood in the genus. This study represents the first broad-scale phylogenetic study of Faramea, with 80 of the c. 170 species sampled, 24 by two or more specimens. We aimed to include specimens representing the entire geographical, morphological and ecological ranges of the genus. Morphological characters historically utilized to delimit infrageneric sections in Faramea (e.g. bract and pyrene forms) were also evaluated. Only one of the currently accepted infrageneric sections was recovered as monophyletic (within a complex of species from other sections) and none of the morphological features traditionally utilized to determine infrageneric relationships in the genus was found to be uniquely diagnostic of a larger clade. Some Faramea lineages appear to be geographically isolated, with several clades containing solely specimens collected in the Atlantic Forest biomes. Of the 24 species represented by at least two specimens, 11 were supported as monophyletic, ten as non-monophyletic and three were not resolved as either monophyletic nor non-monophyletic. The results of the present study constitute a good basis for future studies of taxonomy, biogeography and ecology of Faramea.
The Late Oligocene to Early Miocene flora of the Ban Pa Kha Subbasin (Li Basin, northern Thailand) provides a record of montane dry tropical oak-pine forests. The rich ensemble of Fagaceae typical of these forests might have existed in the wider region of Southeast Asia since Eocene times and various fossil plant assemblages represented both lowland (Fagaceae, Dipterocarpaceae) and upland (Fagaceae, Pinaceae) tropical forests. These findings are in conflict with previous interpretations of vegetation development in northern Thailand, which stressed that stratigraphically older (possibly Late Oligocene) spore and pollen assemblages in northern Thailand were markedly different from the modern tropical flora and had a distinct northern temperate character. A major change in climate would have caused a dramatic shift to tropical conditions since the Mid-Miocene. Considering palaeobotanical data from adjacent regions in Southeast Asia, we suggest that differences in spore and pollen assemblages in intermontane basins in northern Thailand are more likely to represent different facies and lowland/upland settings. Assembly of these forest ecosystems, typically comprising Quercus sections Cyclobalanopsis, Ilex and Quercus, pollen of Castaneoideae with affinities to Castanopsis and Lithocarpus, and extinct fagaceous genera such as Eotrigonobalanus, most probably dates back to the Eocene. The absence of oaks of Quercus section Cerris in the spore and pollen assemblage of the Ban Pa Kha Subbasin, despite this group being part of the modern vegetation, might reflect the late arrival (secondary radiation) of this chiefly temperate group in tropical Southeast Asia.
Pycnandra (Sapotaceae), the largest endemic genus in New Caledonia, comprises 66 species classified in six subgenera. We tested phylogenetic relationships and a proposed infrageneric classification by sampling 60 species for sequences of nuclear ribosomal DNA (ETS, ITS, RPB2) and plastid DNA (trnH–psbA) and nine morphological characters. Data were analysed with Bayesian inference, parsimony jackknifing and lineage through time. We recovered a phylogenetic tree supporting the recognition of six proposed subgenera (Achradotypus, Leptostylis, Pycnandra, Sebertia, Trouettia and Wagapensia). Because a subgeneric classification is used, the nomenclature will be stable when the members are transferred to Pycnandra. Morphological traits were optimized in the BEAST analysis, adding evidence to earlier work that morphology has limited value for successfully diagnosing groups in Sapotaceae. We confirm a previously suspected case of cryptic species that exhibit the same morphological features and require the same abiotic conditions, but are distantly related in the phylogenetic tree. We detected two possible new cases of cryptic sibling species that might warrant recognition. A slowdown in speciation rate in several genera has been suggested as evidence that New Caledonia was once submerged after rifting from Australia. Plotting lineages through time reveals two important intervals at 7.5–8.6 Ma and present to 1.5 Ma, when net molecular diversification within the genus was zero. This indicates that the genus presently has reached a dynamic equilibrium, providing additional evidence that New Caledonia is an old Darwinian island, being submerged during the Eocene and colonized after re-emergence c. 37 Ma.