Scandiceae subtribe Daucinae encompasses umbellifers that have fruits with prominent secondary ridges projecting into wings (former tribe Laserpitieae) or spines (former tribe Caucalideae pro parte). It comprises several economically or medicinally important genera including Cuminum, Daucus, Laser, Laserpitium and Thapsia among others. Recent molecular studies, based mostly on nrDNA ITS sequences, revealed that neither Daucus nor Laserpitium are monophyletic. To address issues of relationships and apply respective nomenclatural changes, we obtained additional ITS sequences as well as independent data from three plastid markers—rps16 intron, rpoC1 intron and rpoB‐trnC intergenic spacer—for a comprehensive sample of the subtribe. We examined data for 260 accessions representing all genera of Daucinae and 81 of its ca. 93 species. Phylogenetic trees were estimated using maximum likelihood and Bayesian inference methods. The results indicate that former Laserpitieae constitute a paraphyletic grade at the base of the spiny‐fruited members of Daucinae while traditionally delimited Daucus and Laserpitium are polyphyletic. To maintain a monophyletic Daucus, we suggest including the following genera and species into its synonymy: Agrocharis, Melanoselinum, Monizia, Pachyctenium, Pseudorlaya, Rouya, Tornabenea, Athamanta dellacellae and Cryptotaenia elegans. The species of Laserpitium occur in seven clades and only six species of the Laserpitium s.str. clade retain the generic name. Several species are transferred to Ekimia, Laser and Thapsia; additionally, a monospecific genus Siler is restored and a new genus, Silphiodaucus, is established. The inclusion of Ammodaucus into Thapsia suggested in an earlier study is not supported. The position of Laserpitium pseudomeum requires further study.
Apiaceae tribe Scandiceae includes species with diverse fruits that depending upon their morphology are dispersed by gravity, carried away by wind, or transported attached to animal fur or feathers. This diversity is particularly evident in Scandiceae subtribe Daucinae, a group encompassing species with wings or spines developing on fruit secondary ribs. In this paper, we explore fruit evolution in 86 representatives of Scandiceae and outgroups to assess adaptive shifts related to the evolutionary switch between anemochory and epizoochory and to identify possible dispersal syndromes, i.e., patterns of covariation of morphological and life-history traits that are associated with a particular vector. We also assess the phylogenetic signal in fruit traits. Principal component analysis of 16 quantitative fruit characters and of plant height did not clearly separate species having different dispersal strategies as estimated based on fruit appendages. Only presumed anemochory was weakly associated with plant height and the flattening of mericarps with their accompanying anatomical changes. We conclude that in Scandiceae, there are no distinct dispersal syndromes, but a continuum of fruit morphologies relying on different dispersal vectors. Phylogenetic mapping of ten discrete fruit characters on trees inferred by nrDNA ITS and cpDNA sequence data revealed that all are homoplastic and of limited use for the delimitation of genera. Spines evolved from wings developing on secondary ribs. We hypothesize that spines cannot form on primary ribs because these contain vascular bundles that may constrain such a transformation. We describe a new subtribe for Artedia and propose three new combinations in Daucus.
Premise Despite intensive research, the pathways and driving forces behind the evolution of derived woodiness on oceanic islands remain obscure. The genus Daucus comprises mostly herbs (therophytes, hemicryptophytes) with few rosette treelets (chamaephytes) endemic to various Macaronesian archipelagos, suggesting their independent evolution. To elucidate the evolutionary pathways to derived woodiness, we examined phylogenetic relationships and the habit and secondary xylem evolution in Daucus and related taxa. Methods Sixty taxa were surveyed for molecular markers, life history, and habit traits. Twenty‐one species were considered for wood anatomical characters. A dated phylogeny was estimated using Bayesian methods. The evolution of selected traits was reconstructed using parsimony and maximum likelihood. Results Daucus dispersed independently to the Canary Islands (and subsequently to Madeira), Cape Verde, and the Azores in the late Miocene and Pleistocene. Life span, reproductive strategy, and life form were highly homoplastic; the ancestor of Daucus was probably a monocarpic, biennial hemicryptophyte. Rosette treelets evolved independently in the Canarian‐Madeiran lineage and in Cape Verde, the latter within the last 0.13 Myr. Treelets and hemicryptophytes did not differ in wood anatomy. Pervasive axial parenchyma in wood occurred more often in polycarpic rather than monocarpic species. Conclusions Life span and life form in Daucus are evolutionarily labile and may change independently of wood anatomy, which is related to plant reproductive strategy rather than to life form. Insular woodiness may evolve rapidly (as demonstrated in D. bischoffii), and in Daucus, it does not seem to be an adaptation to lower the risk of xylem embolism.
Premise Pseudanthia are widespread and have long been postulated to be a key innovation responsible for some of the angiosperm radiations. The aim of our study was to analyze macroevolutionary patterns of these flower‐like inflorescences and their potential correlation with diversification rates in Apiaceae subfamily Apioideae. In particular, we were interested to investigate evolvability of pseudanthia and evaluate their potential association with changes in the size of floral display. Methods The framework for our analyses consisted of a time‐calibrated phylogeny of 1734 representatives of Apioideae and a morphological matrix of inflorescence traits encoded for 847 species. Macroevolutionary patterns in pseudanthia were inferred using Markov models of discrete character evolution and stochastic character mapping, and a principal component analysis was used to visualize correlations in inflorescence architecture. The interdependence between net diversification rates and the occurrence of pseudocorollas was analyzed with trait‐independent and trait‐dependent approaches. Results Pseudanthia evolved in 10 major clades of Apioideae with at least 36 independent origins and 46 reversals. The morphospace analysis recovered differences in color and compactness between floral and hyperfloral pseudanthia. A correlation between pseudocorollas and size of inflorescence was also strongly supported. Contrary to our predictions, pseudanthia are not responsible for variation in diversification rates identified in this subfamily. Conclusions Our results suggest that pseudocorollas evolve as an answer to the trade‐off between enlargement of floral display and costs associated with production of additional flowers. The high evolvability and architectural differences in apioid pseudanthia may be explained on the basis of adaptive wandering and evolutionary developmental biology.
In this monograph, for the first time, the pollen morphology was analysed in the context of modern taxonomic treatment of the order and statistically evaluated in search of traits that could be utilised in further taxonomic and evolutionary studies. Our research included pollen sampled from 417 herbarium specimens representing 158 species belonging to 125 genera distributed among all major lineages of Apiales. The pollen was mechanically isolated, acetolysed, suspended in pure glycerine and mounted on paraffin-sealed slides for light microscopy investigation. Although most of the analysed traits were highly homoplastic and showed significant overlap even between distantly related lineages, we were able to construct a taxonomic key based on characters that bear the strongest phylogenetic signal: P/E ratio, mesocolpium shape observed in polar view and ectocolpus length relative to polar diameter. All the investigated traits are easy to observe with light microscopy and defined by clear and well-documented typology. Early diverging lineages of Apiales constitute a distinct group due to subprolate pollen grains (P/E ratio < 1.25). Among four subfamilies of Apiaceae, Mackinlayoideae can be easily identified based on a combination of traits shared with Klotzschia and Platysace—enigmatic umbllifers with highly uncertain phylogenetic position. Pollen of Azorelloideae is much more diverse but retains many plesiomorphic traits found in early-diverging Apioideae. In contrast, Saniculoideae and most representatives of Apioideae are characterised by evolutionary advanced morphology (perprolate pollen grains with relatively short ectocolpus and bone-shaped outline in colpus view). However, it remains unclear whether similarities between Saniculoideae and higher apioids are an example of convergent evolution or reflect common ancestry. Pollen of Hermas shows a unique combination of traits some of which are typical for Azorelloideae while others resemble Saniculoideae.
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