Phylogenetic placement of the monotypic Baolia (Amaranthaceae s.l.) based on morphological and molecular evidence.

BACKGROUND: Baolia H.W.Kung & G.L.Chu is a monotypic genus only known in Diebu County, Gansu Province, China. Its systematic position is contradictory, and its morphoanatomical characters deviate from all other Chenopodiaceae. Recent study has regarded Baolia as a sister group to Corispermoideae. We therefore sequenced and compared the chloroplast genomes of this species, and resolved its phylogenetic position based on both chloroplast genomes and marker sequences. RESULTS: We sequenced 18 chloroplast genomes of 16 samples from two populations of Baolia bracteata and two Corispermum species. These genomes of Baolia ranged in size from 152,499 to 152,508 bp. Simple sequence repeats (SSRs) were primarily located in the LSC region of Baolia chloroplast genomes, and most of them consisted of single nucleotide A/T repeat sequences. Notably, there were differences in the types and numbers of SSRs between the two populations of B. bracteata. Our phylogenetic analysis, based on both complete chloroplast genomes from 33 species and a combination of three markers (ITS, rbcL, and matK) from 91 species, revealed that Baolia and Corispermoideae (Agriophyllum, Anthochlamys, and Corispermum) form a well-supported clade and sister to Acroglochin. According to our molecular dating results, a major divergence event between Acroglochin, Baolia, and Corispermeae occurred during the Middle Eocene, approximately 44.49 mya. Ancestral state reconstruction analysis showed that Baolia exhibited symplesiomorphies with those found in core Corispermoideae characteristics including pericarp and seed coat. CONCLUSIONS: Comparing the chloroplast genomes of B. bracteata with those of eleven typical Chenopodioideae and Corispermoideae species, we observed a high overall similarity and a one notable noteworthy case of inversion of approximately 3,100 bp. of DNA segments only in two Atriplex and four Chenopodium species. We suggest that Corispermoideae should be considered in a broader sense, it includes Corispermeae (core Corispermoideae: Agriophyllum, Anthochlamys, and Corispermum), as well as two new monotypic tribes, Acroglochineae (Acroglochin) and Baolieae (Baolia).

Protection against insect predation during fruit development: the role of fleshy fruit wings of three species of Zygophyllum in the cold desert of Central Asia.

Introduction: Fruit wings serve various ecological functions, including facilitating wind dispersal, providing physical protection to seeds, and regulating seed germination. While many studies have reported the role of fruit wings in plants, little is known about their protective function during fruit development. Methods: In this study, winged fruits damaged by insects in natural populations of three Zygophyllum species (Z. potaninii, Z. lehmannianum and Z. macropterum) were investigated. We measured and compared the percentage of damaged winged fruits, seed set, seed mass, seed germination, and seedling growth of different insect herbivory categories. Results: The results revealed that the percentage of winged fruits with damaged wings only (low predation) was significantly higher than that of with damaged both fruit wings and fruit bodies (high predation). Furthermore, winged fruits with low predation had significant higher seed set, seed mass, seed germination, and dry mass and relative growth rate (RGR) in the seedlings which grown from the seeds, than that from winged fruits with high predation. Discussion: These results demonstrate that the presence of the fruit wings may provide protection for the seeds to alleviate harm from insect predation before dispersal. These findings provide new insights into the function of fruit wings and the reproductive strategies of desert plants.

Inflated Ovary May Increase the Dispersal Ability of Three Species in the Cold Deserts of Central Asia.

Among the diaspores of angiosperms an inflated ovary (IO) is a novel morphological trait, but no studies have evaluated its effects on dispersal. The primary aim of this study was to determine the effect of the IO on diaspore dispersal in three cold desert species (Carex physodes, Calligonum junceum, and Sphaerophysa salsula). Various morphological features and the mass of fruits and seeds of each species were measured. The role of an IO in diaspore dispersal by wind and water was determined by comparing responses of intact (inflated) IOs and flattened fruits and seeds. Mature diaspores of three species were dispersed by wind, and the IO significantly increased dispersal distance in the field and at different wind speeds in the laboratory. The floating time on water was greater for inflated fruits than flattened fruits and seeds. Since the seed remains inside the IO until after dispersal is completed, the IO of the three species enhances diaspore dispersal. This is the first detailed study on how an IO increases diaspore/seed dispersal. Furthermore, after primary dispersal by wind, secondary dispersal can occur via wind or surface runoff of water, and each method is enhanced by the presence of an IO.

Stamen curvature and temporal flower closure assure reproductive success in an early-spring-flowering perennial in the cold desert of Middle Asia.

Floral organ movements that ensure autonomous selfing are likely to occur in species that grow in habitats with pollinator scarcity and/or an unpredictable environment. Stamen curvature and temporal flower closure are two important floral behaviors that can influence plant pollination mode and reproductive success. However, both behaviors are rarely reported within a species, and little is known about how these two movements of floral organs ensure reproductive success in an unpredictable early spring environment with few pollinators. The aim of this study was to assess whether stamen curvature and temporal flower closure ensure successful reproduction of Leontice incerta in its cold desert habitat. Flowering phenology, floral traits, stamen curvature patterns and flower visitors were surveyed. The breeding system, capacity and timing for autonomous selfing were estimated by pollination manipulations. The timing of floral opening and closure, and benefits of temporal flower closure were determined. We found that flowering of L. incerta began in late March to early April in two populations in two years, and the yellow flowers had neither nectar nor scent. Floral visitation occurred very rarely, but bees (Colletes sp.) were potential pollinators. Fruit and seed set of open and bagged flowers did not differ significantly from that of self-pollinated or cross-pollinated flowers. However, removal of stamens significantly decreased seed set. Self-pollination occurs when the stamens curve and anthers touch the stigma autonomously, suggesting autonomous selfing assurance of seed production in this self-compatible species. Both fruit and seed set of flowers that were prevented from closing were significantly lower than those of control flowers and closed flowers treated with simulated rain treatment. Therefore, stamen curvature and temporal floral closure can ensure successful sexual reproduction of L. incerta in early spring in the cold desert, where lack of pollinators otherwise may lead to pollination failure.

Flexibility of resource allocation in a hermaphroditic-gynomonoecious herb through deployment of female and male resources in perfect flowers.

PREMISE OF THE STUDY: It has been hypothesized that two flower types permit flexible allocation of resources to female and male functions, yet empirical evidence for the sex-allocation hypothesis remains scarce in gynomonoecious species. To characterize resource allocation to pistillate and perfect flowers and allocation of perfect flowers between gynomonoecious and hermaphroditic individuals, we examined the flexibility and whether female-biased allocation increases with plant size in the hermaphroditic-gynomonoecious herb Eremurus anisopterus. METHODS: Frequency of gynomonoecious individuals, flower production, and plant size were investigated in different populations. Floral allocation was compared among the three flower types of E. anisopterus. KEY RESULTS: Frequency of gynomonoecious plants varied from 2-17% in nine populations. Only larger plants produced female flowers at the bottom of racemes. Both female and perfect flower production tended to increase proportionately with plant size in gynomonoecious individuals. Female flowers did not produce less biomass than perfect flowers from hermaphroditic or gynomonoecious plants. However, both female and perfect flowers from gynomonoecious individuals had lighter stamen mass, but larger pistil mass, than perfect flowers from hermaphrodites. CONCLUSIONS: Although the prediction of an increase in female flower number with plant size was not observed in E. anisopterus, the flexibility of sex allocation in gynomonoecious species was confirmed in that gynomonoecious individuals had a female-biased floral allocation compared to hermaphroditic individuals. Such comparisons of gynomonoecious to hermaphroditic individuals permit us to unveil a sexual adjustment strategy: flexibility of sexual investments within plants.

Intermediate complex morphophysiological dormancy in seeds of the cold desert sand dune geophyte Eremurus anisopterus (Xanthorrhoeaceae; Liliaceae s.l.).

BACKGROUND AND AIMS: Little is known about morphological (MD) or morphophysiological (MPD) dormancy in cold desert species and in particular those in Liliaceae sensu lato, an important floristic element in the cold deserts of Central Asia with underdeveloped embyos. The primary aim of this study was to determine if seeds of the cold desert liliaceous perennial ephemeral Eremurus anisopterus has MD or MPD, and, if it is MPD, then at what level. METHODS: Embryo growth and germination was monitored in seeds subjected to natural and simulated natural temperature regimes and the effects of after-ripening and GA3 on dormancy break were tested. In addition, the temperature requirements for embryo growth and dormancy break were investigated. KEY RESULTS: At the time of seed dispersal in summer, the embryo length:seed length (E:S) ratio was 0·73, but it increased to 0·87 before germination. Fresh seeds did not germinate during 1 month of incubation in either light or darkness over a range of temperatures. Thus, seeds have MPD, and, after >12 weeks incubation at 5/2 °C, both embryo growth and germination occurred, showing that they have a complex level of MPD. Since both after-ripening and GA3 increase the germination percentage, seeds have intermediate complex MPD. CONCLUSIONS: Embryos in after-ripened seeds of E. anisopterus can grow at low temperatures in late autumn, but if the soil is dry in autumn then growth is delayed until snowmelt wets the soil in early spring. The ecological advantage of embryo growth phenology is that seeds can germinate at a time (spring) when sand moisture conditions in the desert are suitable for seedling establishment.

Pistillate flowers experience more pollen limitation and less geitonogamy than perfect flowers in a gynomonoecious herb.

Gynomonoecy, a sexual system in which plants have both pistillate (female) flowers and perfect (hermaphroditic) flowers, occurs in at least 15 families, but the differential reproductive strategies of the two flower morphs within one individual remain unclear. Racemes of Eremurus anisopterus (Xanthorrhoeaceae) have basal pistillate and distal perfect flowers. To compare sex allocation and reproductive success between the two flower morphs, we measured floral traits, pollinator preferences, and pollen movement in the field. Pollen limitation was more severe in pistillate flowers; bee pollinators preferred to visit perfect flowers, which were also capable of partial self-fertilization. Pollen-staining experiments indicated that perfect flowers received a higher proportion of intra-plant pollen (geitonogamy) than pistillate flowers. Plants with greater numbers of pistillate flowers received more outcross pollen. The differential reproductive success conformed with differential floral sex allocation, in which pistillate flowers produce fewer but larger ovules, resulting in outcrossed seeds. Our flower manipulations in these nectarless gynomonoecious plants demonstrated that perfect flowers promote seed quantity in that they are more attractive to pollinators, while pistillate flowers compensate for the loss of male function through better seed quality. These results are consistent with the outcrossing-benefit hypothesis for gynomonoecy.