Pseudodiplostemony, and its implications for the evolution of the androecium in the Caryophyllaceae

The androecium of the Caryophyllaceae is varied, ranging from a two-whorled condition to a single stamen. A number of species belonging to the three subfamilies, Caryophyl-loideae, Alsinoideae and Paronychioideae have been studied ontogenetically with the SEM to understand their peculiar androecial development in the broader context of the Caryophyllales alliance. Although patterns of initiation are highly variable among species, there are three ontogenetic modes of stamen initiation: all stamens simultaneous within a whorl, the antepetalous stamens simultaneous and the antesepalous sequentially with a reversed direction, or both whorls sequentially with or without a reversed direction. The most common floral (ontogenetic) sequence of the Caryophyllaceae runs as follows: five sepals (in a 2/5 sequence), the stamens in front of the three inner sepals successively, stamens opposite the two outermost sepals, five antepetalous stamens (simultaneously or in a reversed spiral superimposed on the spiral of the antesepalous stamens), five outer sterile (petaloid) organs arising before, simultaneously or after the antesepalous stamens, often by the division of common primordia. A comparison with the floral configurations of the Phytolaccaceae and Molluginaceae indicates that the outer petaline whorl of the Caryophyllaceae corresponds positionally to the alternisepalous stamens of somePhytolacca, such asP. dodecandra. The difference withP. dodecandra lies in the fact that an extra inner or outer whorl is formed in the Caryophyl-laceae, in alternation with the sepals. A comparable arrangement exists in the Molluginaceae, though the initiation of stamens is centrifugal. A comparison of floral ontogenies and the presence of reduction series in the Caryophyllaceae support the idea that the pentamerous arrangement is derived from a trimerous prototype. Petals correspond to sterillized stamens and are comparable to two stamen pairs opposite the outer sepals and a single stamen alternating with the third and fifth sepals. Petals are often in a state of reduction; they may be confused with staminodes and they often arise from common stamenpetal primordia. The antesepalous stamen whorl represents an amalgamation of two whorls: initiation is reversed with the stamens opposite the fourth and fifth formed sepals arising before the other, while the stamens opposite the first and second formed sepals are frequently reduced or lost. Reductive trends are correlated with the mode of initiation of the androecium, as well as changes in the number of carpels, and affect the antesepalous and antepetalous whorls in different proportions. It is concluded that the androecium of the Caryophyllaceae is pseudodiplos-temonous and is not comparable to diplostemonous forms in the Dilleniidae and Rosidae. The basic floral formula of Caryophyllaceae is as follows: sepals 5—petals 5 (sterile stamens)—antesepalous stamens 3+2—antepetalous stamens 5 gynoecium 5.     

Delimitation and phylogeny of Aletris (Nartheciaceae) with implications for perianth evolution

Aletris,containing approximately 21 species,is the largest genus in Nartheciaceae,and is disjunctively distributed in eastern Asia and eastern North America.Its delimitation has been controversial beca...     

The discovery of polyandry in Curculigo (Hypoxidaceae): implications for androecium evolution of asparagoid monocotyledons

BACKGROUND AND AIMS: Individual flowers of the monocot Curculigo racemosa (Hypoxidaceae, Asparagales) are regularly polyandrous. To evaluate the significance of this almost unique character among Asparagales for flower evolution of asparagoid monocots, flowers of C. racemosa were studied comparatively. METHODS: Anthetic flowers as well as early floral developmental stages were studied by light and scanning electron microscopy. KEY RESULTS: Despite the polyandry, floral development is similar to that of other Asparagales with a developmental gradient from adaxial to abaxial. Stamens initiate simultaneously and the diameter of staminal primordia is about half of that in species with six anthers. The number of stamens is not fixed (12-26) and varies within the same inflorescence. Surprisingly, the gynoecium can be four- or six-locular, besides the normal trimerous state. CONCLUSIONS: The discovery of a polyandrous Curculigo reveals plasticity of stamen number at the base of Asparagales. Orchidaceae - sister to all other Asparagales - has a reduced stamen number (three, two or one), whereas in Hypoxidaceae - part of the next diverging clade - either the normal monocot stamen number (six), polyandry (this study) or the loss of three anthers (Pauridia) occurs. However, at present it is impossible to decide whether the flexibility in stamen number is autapomorphic for each group or whether it is a synapomorphy. The small size of stamen primordia of Curculigo is conspicuous. It allows more space for additional androecial primordia. Stamens are initiated as independent organs, and filaments are not in bundles, hence C. racemosa is not secondarily polyandrous as may be the case in the distantly related Gethyllis of asparagoid Amaryllidaceae. The increase in carpel number is a rare phenomenon in angiosperms. A possible explanation for the polyandry of C. racemosa is that it is a natural SUPERMAN-deficient mutant, which shows an increase of stamens, or ULTRAPETALA or CARPEL FACTORY mutants, which are polyandrous and changed in carpel number.     

Structure and evolution of the androecium in the Malvatheca clade (Malvaceae s.l.) and implications for Malvaceae and Malvales

Androecial development and structure as well as floral vasculature of six selected species of Bombacoideae and of several smaller lineages of the Malvatheca clade (Malvaceae s.l.) were studied. All studied taxa share a similar pattern of androecial development: initially, five antepetalous/antetepalous and five alternipetalous/alternitepalous primary androecial primordia develop on a ring wall. Two elongate secondary androecial primordia form on each antepetalous/antetepalous sector. At anthesis the androecium consists of an androecial tube crowned by five androecial lobes. Each of these lobes is the developmental product of an alternipetalous/alternitepalous primary androecial primordium and its two neighbouring antepetalous/antetepalous secondary androecial primordia. The elongate, sessile androecial units are positioned along the lateral margins of the androecial lobes and in the distal part of the androecial tube. Seen in the light of the most recent studies of floral development and phylogeny of the Malvaceae and the Malvales as a whole, our data indicate that i) elongate, sessile androecial units are ancestral in the Malvatheca clade, that ii) an obdiplostemonous floral ground plan is a synapomorphy for the Malvaceae, and that iii) diplostemony is most likely ancestral in the Malvales.     

Caryophyllales: a key group for understanding wood anatomy character states and their evolution

Definitions of character states in woods are softer than generally assumed, and more complex for workers to interpret. Only by a constant effort to transcend the limitations of glossaries can a more than partial understanding of wood anatomy and its evolution be achieved. The need for such an effort is most evident in a major group with sufficient wood diversity to demonstrate numerous problems in wood anatomical features. Caryophyllales s.l., with approximately 12 000 species, are such a group. Paradoxically, Caryophyllales offer many more interpretive problems than other ‘typically woody’ eudicot clades of comparable size: a wider range of wood structural patterns is represented in the order. An account of character expression diversity is presented for major wood characters of Caryophyllales. These characters include successive cambia (more extensively represented in Caryophyllales than elsewhere in angiosperms); vessel element perforation plates (non‐bordered and bordered, with and without constrictions); lateral wall pitting of vessels (notably pseudoscalariform patterns); vesturing and sculpturing on vessel walls; grouping of vessels; nature of tracheids and fibre‐tracheids, storying in libriform fibres, types of axial parenchyma, ray anatomy and shifts in ray ontogeny; juvenilism in rays; raylessness; occurrence of idioblasts; occurrence of a new cell type (ancistrocladan cells); correlations of raylessness with scattered bundle occurrence and other anatomical discoveries newly described and/or understood through the use of scanning electron microscopy and light microscopy. This study goes beyond summarizing or reportage and attempts interpretations in terms of shifts in degrees of juvenilism, diversification in habit, ecological occupancy strategies (with special attention to succulence) and phylogenetic change. Phylogenetic change in wood anatomy is held to be best interpreted when accompanied by an understanding of wood ontogeny, species ecology, species habit and taxonomic context. Wood anatomy of Caryophyllales demonstrates problems inherent in binary character definitions, mapping of morphological characters onto DNA‐based trees and attempts to analyse wood structure without taking into account ecological and habital features. The difficulties of bridging wood anatomy with physiology and ecology are briefly reviewed. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 342–393.     

Female flower and fruit anatomy of Tetroncium magellanicum: implications for gynoecium evolution in the early divergent monocot order Alismatales

Female flower and fruit anatomy, including vasculature, are studied for the first time in Tetroncium (Juncaginaceae: Alismatales). Other members of Juncaginaceae (and the relatively close Maundiaceae) possess a peculiar type of gynoecium with pronounced carpel fusion via the floral centre. Their carpels are supplied by individual vascular traces and can be interpreted either as synascidiate (if viewed as horizontally inserted) or free and plicate (if viewed as obliquely inserted on an elongated receptacle). In Tetroncium, the gynoecium is tetracarpellate and clearly has a well‐developed synascidiate zone with septa formed by united flanks of adjacent carpels. The gynoecium of Tetroncium is supplied by a common ring of vascular tissue that splits into dorsal and heterocarpellary ventral (synventral) bundles, a condition that can be expected in a typical syncarpous gynoecium. The fruit is indehiscent and contains one or two seeds. The syncarpy of Tetroncium is of functional significance for fruit formation, as it allows the thin septa to be distorted, thus providing more space for the developing seed(s). The occurrence of typical syncarpy in Tetroncium provides further evidence for the highly homoplastic evolution of gynoecium characters in the early‐divergent monocot order Alismatales. Either the similarity between gynoecia of Maundiaceae and Triglochin (Juncaginaceae) is due to parallel evolution or the syncarpy of Tetroncium should be viewed as secondarily derived. In the latter scenario, fusion via the floral centre is probably a synapomorphy of core Alismatales (Helobiae) and more typical syncarpy evolved independently in several lineages, such as Scheuchzeria, Tetroncium and Butomus/Hydrocharitaceae. In total, Tetroncium differs from other Juncaginaceae in 13 structural characters, including ensiform leaves that are similar to those of Tofieldiaceae. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 712–724.     

Delimitation and phylogeny of Aletris (Nartheciaceae) with implication for perianth evolution

Aletris, containing approximately 21 species, is the largest genus in Nartheciaceae, and is disjunctively distributed in eastern Asia and eastern North America. Its delimitation has been controversial because it is uncertain whether Metanarthecium should be included in the genus. Although there are a few molecular phylogenetic studies on Aletris, the interspecific relationships within the genus have never been evaluated in a phylogenetic context. Here we used two cpDNA loci, matK and trnL-F, to delimitate Aletris and discuss the phylogeny within the genus. Phylogenetic analyses showed Metanarthecium might be distantly related to Aletris. This is also supported by morphological, palynological, cytological, and phytochemical data. Therefore, Metanarthecium should be excluded from Aletris. Within Aletris, there are two major clades: A. farinosa and A. lutea of eastern North America and A. glabra of eastern Asia form clade A; and the remaining Asian species form clade B. The Asian clade includes three subclades: subclade I (two varieties of A. pauciflora, and A. glandulifera and A. megalantha), subclade II (three samples of A. laxiflora), and subclade III (all other sampled Asian species). Based on phylogenetic relationships, A. pauciflora var. khasiana deserves a specific status, and A. gracilipes, formerly a synonym of A. laxiflora, should be reinstated. The reconstruction of the perianth evolution indicates that perianth connate halfway and glabrous on abaxial surface are plesiomorphic for Aletris and Nartheciaceae. Farinose-glutinous perianth is a diagnostic character for clade A.     

Molecular systematics of the Catesbaeeae-Chiococceae complex (Rubiaceae): flower and fruit evolution and biogeographic implications

The classification of the Catesbaeeae and Chiococceae tribes, along with that of the entire Rubiaceae, has long been debated. The Catesbaeeae-Chiococceae complex (CCC) includes approximately 28 genera and 190 species primarily concentrated in the Greater Antilles (nearly 70% of the species), Central and South America, and in the western Pacific (three genera). Previous molecular studies, with broad sampling of the Rubiaceae, have shown the CCC to be a monophyletic group. The present study is a more detailed examination of the generic relationships within the CCC using two data sets, the nuclear ribosomal ITS regions and the trnL-F chloroplast intron and spacer. Maximum parsimony analyses lend further support to the previous hypotheses that the CCC is monophyletic and sister to Strumpfia maritima. However, within the complex several genera do not form monophyletic groups. Previous studies of the Rubiaceae suggest that the ancestral fruit type in the CCC is a multiseeded capsule. Indehiscent, fleshy fruits appear to have evolved three to four times within this lineage. Changes in floral morphologies within the complex tend to correspond to cladogenesis among and within genera. Finally, molecular analyses suggest one or possibly two long-distance dispersals from the Americas to the western Pacific.     

More is better: the uses of developmental genetic data to reconstruct perianth evolution

The origin and evolution of the perianth remains enigmatic. While it seems likely that an undifferentiated perianth consisting of tepals arose early in angiosperm evolution, it is unclear when and how differentiated perianths consisting of distinct organs, such as petals and sepals, arose. Phylogenetic reconstructions of ancestral perianth states across angiosperms have traditionally relied on morphological data from extant species, but these analyses often produce equivocal results. Here we describe the use of developmental genetic data as an additional strategy to infer the ancestral perianth character state for different angiosperm clades. By assessing functional data in combination with expression data in a maximum likelihood framework, we provide a novel approach for investigating the evolutionary history of the perianth. Results of this analysis provide new insights into perianth evolution and provide a proof of concept for using this strategy to explore the incorporation of developmental genetic data in character state reconstructions. As the assumptions outlined here are tested and more genetic data are generated, we hope that ancestral state reconstructions based on multiple lines of evidence will converge.     

Phylogenetic relationships and taxonomic ranking of pipizine flower flies (Diptera: Syrphidae) with implications for the evolution of aphidophagy

The taxonomic rank and phylogenetic relationships of the pipizine flower flies (Diptera: Syrphidae: Pipizini) were estimated based on DNA sequence data from three gene regions (COI, 28S and 18S) and 111 adult morphological characters. Pipizini has been treated as a member of the subfamily Eristalinae based on diagnostic adult morphological characteristics, while the larval feeding mode and morphology is shared with members of the subfamily Syrphinae. We analysed each dataset, both separately and combined, in a total evidence approach under maximum parsimony and maximum likelihood. To evaluate the influence of different alignment strategies of rDNA 28S and 18S genes on the resulting topologies, we compared the topologies inferred from a multiple alignment using fast Fourier transform (MAFFT) program with those topologies resulting from aligning the secondary structure of these rDNA genes. Total evidence analyses resolved pipizines as a sister group of the subfamily Syrphinae. Although the structural alignment and the MAFFT alignment differed in the inferred relationships of some clades and taxa, there was congruence in the placement of pipizines. The homogeneous morphology of the Pipizini clade in combination with their unique combination of characters among the Syrphidae suggest a change of rank to subfamily. Thus, we propose to divide Syrphidae into four subfamilies, including the subfamily Pipizinae stat. rev.     

Flora of Nigeria: Caryophyllales

Roupala barnettiae from the Monta?as de Misisí in Edo. Trujillo, Venezuela is described and illustrated. Its taxonomic affinities are also discussed.     

Patterns of ontogenetic evolution in perianth diversification of Besseya (Scrophulariaceae)

Species of Besseya have perianth diversity centered largely in meristic variation, extreme corolla diminution, and corolla tube loss. Flowers of Besseya differ from those of their closest relative Synthyris in having a bilabiate corolla. The avenues of ontogenetic evolution that were important in creating these aspects of perianth diversity were explored by optimizing developmental transformations on cladograms. Three different evolutionary transformations were important in the diversification of calyx ontogenies in Besseya, including the derived reciprocal substitution of ontogenetic states in B. bullii, labile abaxial lobe expression in B. wyomingensis, and a heterotopic novel substitution in B. oblongifolia. The bilabiate corolla of Besseya arose via fractionation and localization of the zonal growth expressed in the more plesiomorphic Synthyris. Besseya plantaginea, B. ritteriana, and B. oblongifolia form a group characterized by diminished expression of the anterior corolla lobe. Four alternative scenarios equally explain the ontogenetic transformations underlying extreme corolla diminution and corolla tube loss. Corolla tube loss has been considered rare or impossible among angiosperms, but some alternative scenarios in Besseya demonstrate how this may have occurred via fractionation of zonal growth. Corolla tube loss may have arisen only indirectly from the sympetalous ancestors of Besseya following extreme corolla diminution and the reestablishment of a “full-size” corolla which had a novel developmental pattern that did not include the distribution of zonal growth present in ancestors.     

Frequency–dependent pollinator foraging in polymorphic Clarkia xantiana ssp. xantiana populations: implications for flower colour evolution and pollinator interactions

Under many circumstances pollinators are expected to practice positive frequency–dependent foraging in colour-polymorphic plant populations. Theory suggests, however, that competition for floral resources might favor negative frequency–dependent foraging by some pollinator species, possibly contributing to the maintenance of flower colour variation by negative frequency–dependent selection. We addressed this idea with pollination studies of the California annual plant Clarkia xantiana ssp. xantiana (Onagraceae), which is polymorphic for the presence of conspicuous petal spots and is pollinated by several specialist bee species. At the level of entire pollinator assemblages, we did not detect significant fixed flower colour preferences or frequency–dependent foraging. Three species of specialist bee pollinators, however, showed contrasting forms of frequency–dependent foraging. The most widespread species, Hesperapis regularis (Melittidae) exhibited positive frequency dependence. Two other common species, Lasiglossum pullilabre (Halictidae) and Ceratina sequoiae (Apidae), preferred to visit whichever morph (unspotted or spotted) was locally in the minority. All three species were found to be effective at transferring C. xantiana pollen; H. regularis appeared most effective. Our findings suggest that a mixture of positive and negative frequency–dependent selection on flower colour occurs in C. xantiana , with the form and intensity of selection varying in space and time with pollinator assemblages. Negative frequency–dependent selection via pollination dynamics may play a larger role in maintaining genetic variation in flower colour than was previously thought. Our results also suggest an unappreciated form of niche partitioning among specialist pollinators. Genetic polymorphism in flower colour may sometimes facilitate pollinator coexistence.     

Adaptive mutation: implications for evolution

Adaptive mutation is defined as a process that, during nonlethal selections, produces mutations that relieve the selective pressure whether or not other, nonselected mutations are also produced. Examples of adaptive mutation or related phenomena have been reported in bacteria and yeast but not yet outside of microorganisms. A decade of research on adaptive mutation has revealed mechanisms that may increase mutation rates under adverse conditions. This article focuses on mechanisms that produce adaptive mutations in one strain of Escherichia coli, FC40. These mechanisms include recombination-induced DNA replication, the placement of genes on a conjugal plasmid, and a transient mutator state. The implications of these various phenomena for adaptive evolution in microorganisms are discussed.     

Cultural evolution: implications for understanding the human language faculty and its evolution

Human language is unique among the communication systems of the natural world: it is socially learned and, as a consequence of its recursively compositional structure, offers open-ended communicative potential. The structure of this communication system can be explained as a consequence of the evolution of the human biological capacity for language or the cultural evolution of language itself. We argue, supported by a formal model, that an explanatory account that involves some role for cultural evolution has profound implications for our understanding of the biological evolution of the language faculty: under a number of reasonable scenarios, cultural evolution can shield the language faculty from selection, such that strongly constraining language-specific learning biases are unlikely to evolve. We therefore argue that language is best seen as a consequence of cultural evolution in populations with a weak and/or domain-general language faculty.