Testing the 'assortative mating' hypothesis on a variation maintenance mechanism for flowering time within a forest-floor population of Primula sieboldii

Flowering phenology and allozyme variation were studied to test the existence of positive assortative mating for flowering time in a natural population of Primula sieboldii E. Morren, a heterostylous perennial herb, consisting of approximately 180 genets in a deciduous forest. There was significant variation in flowering date among genets, but not between heterostylous morphs. The temporal order of the flowering time of genets was fairly constant for the two years of the study. The spatial heterogeneity of light availability at the study site was small during the flowering season of the species. In order to analyze the extent of genetic differentiation between early- and late-flowering genet groups, allozyme diversities were analyzed with 10 loci. The G_ST between the early- and late-flowering groups was not significantly different from zero. Evidence of positive assortative mating for flowering time was not detected. Prolongation of flowering duration due to pollen limitation may be one important factor preventing the genetic differentiation of early- and late-flowering groups by enhancing the overlap of flowering time among genets.     

Ecological consequences of a massive flowering event of bamboo (Chusquea culeou) in a temperate forest of Patagonia,Argentina

Question: What changes occur as a consequence of the massive flowering and senescence of the dominant understory species of bamboo, Chusquea culeou (E. Desvaux)? In this study, we documented some of the ecological consequences of this rare event that occurred in 2001, the previous flowering having occurred more than 60 years ago. Location: Nothofagus temperate forest, Patagonia, Argentina. Methods: We assessed changes in environmental variables and bamboo biomass post‐flowering in an old‐growth southern beech forest. In addition, we monitored the demography of emergent Ch. culeou seedling and Nothofagus nervosa saplings, comparing non‐flowered (live understory) and flowered (senescent understory) patches within the forest matrix. Results: Bamboo flowering dramatically increased light availability in the forest understory but, surprisingly, other environmental changes were not observed. Bamboo seedlings emerged in both patch types, and experienced gradual but modest mortality through time. Bamboo dieback promoted higher survivorship and an increment in biomass, height, number of leaves and buds in the saplings of Nothofagus nervosa. Conclusion: The high density of bamboo seedlings 5 years after the flowering event and the independence of emergence from environmental variables suggest that understory regeneration is a gradual process that is not strongly regulated by initial seedling density or resource limitation. In contrast, microenvironmental conditions created after the flowering event significantly increased Nothofagus sapling growth and survival. These results suggest that overstory forest regeneration could be enhanced in this temperate forest in the first years after this infrequent bamboo flowering event.     

Clonal identification by microsatellite loci in sporadic flowering of a dwarf bamboo species, <Emphasis Type="Italic">Sasa cernua</Emphasis>

Dwarf bamboo species are monocarpic. They flower simultaneously and die after several decades. The type of flowering in the genus Sasa ranges from sporadic to gregarious. In order to determine whether or not the sporadic flowering of dwarf bamboo is fixed genetically, we investigated the distribution of clones using eight microsatellite (SSR) loci in a sporadic flowering patch of Sasa cernua Makino, a major dwarf bamboo species found in central Hokkaido. In May 2006, flowering occurred on 60.5% of living culms in a 1600 m² patch. We established a 50 × 10 m study plot in this patch and noted 1267 clumps consisting of 2529 living culms. We investigated all 1267 clumps and identified six multilocus genotypes as clones using five variable SSR loci. All flowering clumps belonged to the same clone. On the other hand, non-flowering vegetative clumps were also discovered to be of the same clone. These data suggest that all flowering culms originated from a single clone of a sporadic flowering patch of S. cernua. Clonal analysis for investigation of sporadic flowering of S. cernua revealed that only a portion of a clone flowers and dies instead of the whole clone.     

Irregular droughts trigger mass flowering in aseasonal tropical forests in asia

General flowering is a community-wide masting phenomenon, which is thus far documented only in aseasonal tropical forests in Asia. Although the canopy and emergent layers of forests in this region are dominated by species of a single family, Dipterocarpaceae, general flowering involves various plant groups. Studying proximate factors and estimating the flowering patterns of the past and future may aid our understanding of the ecological significance and evolutionary factors behind this phenomenon. Here we show that this phenomenon is most likely triggered by irregular droughts based on 10 years of observations. In the aseasonal forests of SE Asia, droughts tend to occur during transition periods from La Ni?a to El Ni?o, which results in an irregular 6-7-yr cycle involving a dry period with several droughts and a wet period without droughts. The magnitude of a flowering event also depends on the timing of droughts associated with the El Ni?o southern oscillation (ENSO) cycle, with the largest events occurring after an interval of several years with no flowering. Because most plant species can only reproduce successfully during large flowering events, changes in the ENSO cycle resulting from global warming, may have serious ramifications for forest regeneration in this region.     

Clonal Structure,Seed Set,and Self-Pollination Rate in Mass-Flowering Bamboo Species during Off-Year Flowering Events

Bamboos are typical examples of highly synchronized semelparous species. Their mass-flowering events occur at supra-annual intervals but they sometimes flower on a small scale in off-years. If some bamboo ramets (culms) of a genet flower and die in off-years, whereas other culms of the same genet do not flower synchronously, the genet can still survive blooming in an off-year and could participate in the next mass-flowering event. At genet level, the effect might be similar to that achieved by synchronously reproducing iteroparous plants. In addition, if multiple genets flower simultaneously in off-years, cross-pollination will be promoted. However, it is not known whether all the culms in a genet flower synchronously and whether multiple genets flower in off-years. We determined the clonal structure of three temperate dwarf bamboo species, i.e., Sasa senanensis, S. kurilensis, and S. palmata, at 24 off-year flowering sites and the surrounding areas in northern Japan using seven microsatellite markers. We also estimated seed set at seven of the sites and self-pollination rates at five sites to determine off-year reproductive success. Next, we investigated whether seed sets at the culm level were related to flowering area and/or number of flowering genets, using generalized linear mixed-effect models (GLMMs). Multiple genets flowered at 9/24 flowering sites. We found that 40/96 of the genets identified had some flowering culms. Non-flowering culms were present in 24/40 flowering genets. Seed set was in the range 2.2%–12.5% and the self-pollination rate was 96.3%. In the best GLMM, seed set increased with flowering area. Seeds were produced in off-years, but cross-pollination was rare in off-years. We suggest that some dwarf bamboos may exhibit iteroparity or imperfectly synchronized semelparity at the genet level, a characteristic similar to that of other reproductively synchronous plants. We also found synchronous flowering of a few genets even in off-years.     

Changes in first flowering dates and flowering duration of 232 plant species on the island of Guernsey

Climate change has affected plant phenology; increasing temperatures are associated with advancing first flowering dates. The impact on flowering duration, however, has rarely been studied. In this study, we analysed first flowering dates and flowering durations from a 27 year dataset of weekly flower observations on 232 plant species from the island of Guernsey in the English Channel. The aim of this study was to explore variation in trends and relationships between first flowering dates, flowering duration and temperature. We specifically looked for evidence that traits, such as life forms and phylogenetic groups, explained variation in sensitivity of first flowering and flowering duration among species. Overall trends revealed significantly earlier flowering over time, by an average of 5.2 days decade^?1 since 1985. A highly significant shortening of flowering duration was observed by an average of 10 days decade^?1. Correlations between first flowering, flowering duration and year varied between different species, traits and flowering periods. Significant differences among traits were observed for first flowering and to a lesser degree for flowering duration. Overall, in comparison to first flowering, more species had significant trends in flowering duration. Temperature relationships revealed large differences in strength and direction of response. 55% of the species revealed a significant negative relationship of first flowering dates and temperature. In contrast, only 19% of flowering durations had a significant negative temperature relationship. The advance in first flowering date together with a shortening of flowering duration suggests potentially serious impacts on pollinators, which might pose a major threat to biodiversity, agriculture and horticulture. Human health, in terms of pollen allergies, however, might benefit from a shortening of specific plant pollen seasons.     

Genets of dwarf bamboo do not die after one flowering event: evidence from genetic structure and flowering pattern

Dwarf bamboos in the genus Sasa are believed to be long-lived, synchronously flowering, and monocarpic plants. However, the monocarpy of dwarf bamboo has not been confirmed, because whether all ramets within one genet flower at the same time cannot be determined without differentiating the genetic structure among ramets. This study aims to evaluate the reproductive traits of Sasa pubiculmis by verifying the monocarpy and physiological integration between flowering ramets and non-flowering ramets during a 4-year flowering period. One genotypically identified genet, which covered an area of approximately 3 ha, had both flowering and non-flowering patches of ramets during the 4-year flowering period (2004–2007). A fraction of the flowering genet remained non-flowering during the 4 years of observation, and did not die after mass flowering. Flowering ramets were physically connected to non-flowering ramets via rhizomes, and assimilated ¹³C was allocated from non-flowering ramets to flowering ramets. Consequently, we clarified that this dwarf bamboo potentially has polycarpic reproductive traits rather than monocarpic, and a genet can keep rhizomes and non-flowering patches alive to sustain the organism after mass flowering.     

Evolutionary origin of a periodical mass‐flowering plant

The evolutionary origin of periodical mass‐flowering plants (shortly periodical plants), exhibiting periodical mass flowering and death immediately after flowering, has not been demonstrated. Within the genus Strobilanthes (Acanthaceae), which includes more than 50 periodical species, Strobilanthes flexicaulis on Okinawa Island, Japan, flowers gregariously every 6 years. We investigated the life history of S. flexicaulis in other regions and that of closely related species together with their molecular phylogeny to reveal the evolutionary origin of periodical mass flowering. S. flexicaulis on Taiwan Island was found to be a polycarpic perennial with no mass flowering and, in the Yaeyama Islands, Japan, a monocarpic perennial with no mass flowering. Molecular phylogenetic analyses indicated that a polycarpic perennial was the ancestral state in this whole group including S. flexicaulis and the closely related species. No distinctive genetic differentiation was found in S. flexicaulis among all three life histories (polycarpic perennial, monocarpic perennial, and periodical plant). These results suggest that among S. flexicaulis, the periodical mass flowering on Okinawa Island had evolved from the polycarpic perennial on Taiwan Island via the monocarpic perennial in the Yaeyama Islands. Thus, the evolution of life histories could have taken at the level of local populations within a species.     

Discovery of a life history shift: precocious flowering in an introduced population of Prosopis

The genus Prosopis contains many valuable, long-lived, multi-purpose legume trees, some of which are also invasive species. The time of first flowering is important for increasing production of sweet, protein-rich pods in plantations, especially under short rotations, and affects the rate of spread as a weed. Trees generally begin flowering at 3–5 years of age. However, seedlings from a small seedlot collected from an introduced population in southern Mauritania were observed to begin flowering at 3–4 months after germination. This is unknown in the native range of any Prosopis species, and appears to represent a major evolutionary event triggered after naturalization. This paper reports a detailed investigation with seed collected from Aleg, Brakna region, Mauritania, in 1998 (Prosopis sp. ‘Aleg’). Two experiments were established, in glasshouses at Cirad, Montpellier, France, and Coventry University, UK. Flowering began at the two sites 104 and 169 days after sowing, respectively, and 97.5% of plants had flowered at Coventry after 306 days. Flowers produced abundant pollen with 50–60% viability indicated by FDA staining and 40–50% germination on an artificial medium. No pods were formed. All plants analysed were diploid. Morphologically, these juvenile plants were similar to American species of section Algarobia, and exhibited leaf characters typical of P. pallida. However, due to the large variation in morphology within this section of the genus, further studies are required to confirm the actual species. This is the first record of precocious flowering in Prosopis, and has been found only in a single, introduced population. Implications for genetic improvement of tropical Prosopis, and particularly the spread of these species as invasive weeds, are discussed. Such a life history shift in the increased reproductive ability of a species following introduction, with potentially significant environmental effects, may not be restricted to Prosopis and merits further detailed investigation.     

Insect assemblages on flowering patches of 12 bamboo species

Bamboos are known as long-living monocarpic plants that exhibit synchronous flowering at long intervals. It has been reported that florivory has a critical effect on their seed production in bamboos, especially in small scale flowering patches. In this study, we aimed to determine species composition and life history of florivorous insects in multiple bamboo species. We collected the inflorescences of 12 bamboo species from 15 sites in Japan from 2017 to 2019 and recorded insect assemblages found in the flowers. Five different insect species were observed to feed on the flowers of bamboo species. Among these, the larvae of two Dicraeus species were the most widely observed florivorous insects of the bamboo species. The other insects included the larvae of Cecidomyiidae sp., which was frequently observed on Sasa species, and the other two insects were Epuraea submicrurula, and Dimorphopterus japonicus. These insects were oviposited on the florets during the budding period. In addition, predators and parasitoids were found in the bamboo flowers. Our findings suggest that the bamboos flowers were mainly consumed by dipteran larvae. Insect species composition and frequency varied among bamboo species and sites. Comparing the frequency of the florivorous insects among bamboo flowering patches, including mass flowering patches, in different areas is needed in future studies.     

The strength of assortative mating for flowering date and its basis in individual variation in flowering schedule

Although it has been widely asserted that plants mate assortatively by flowering time, there is virtually no published information on the strength or causes of phenological assortment in natural populations. When strong, assortative mating can accelerate the evolution of plant reproductive phenology through its inflationary effect on genetic variance. We estimated potential assortative mating for flowering date in 31 old‐field species in Ontario, Canada. For each species, we constructed a matrix of pairwise mating probabilities from the individual flowering schedules, that is the number of flower deployed on successive dates. The matrix was used to estimate the phenotypic correlation between mates, ρ, for flowering date. We also developed a measure of flowering synchrony within species, S, based upon the eigenstructure of the mating matrix. The mean correlation between pollen recipients and potential donors for flowering date was  = 0.31 (range: 0.05–0.63). A strong potential for assortative mating was found among species with high variance in flowering date, flowering schedules of short duration and skew towards early flower deployment. Flowering synchrony, S, was negatively correlated with potential assortment (r = ?0.49), but we go on to show that although low synchrony is a necessary condition for phenological assortative mating, it may not be sufficient to induce assortment for a given phenological trait. The potential correlation between mates showed no seasonal trend; thus, as climate change imposes selection on phenology through longer growing seasons, spring‐flowering species are no more likely to experience an accelerated evolutionary response than summer species.     

Plant reproductive phenology over four years including an episode of general flowering in a lowland dipterocarp forest,Sarawak, Malaysia

The first systematic observation of a general flowering, a phenomenon unique to lowland mixed-dipterocarp forests in Southeast Asia, is presented. During general flowering, which occurs at irregular intervals of 3–10 yr, nearly all dipterocarp species together with species of other families come heavily into flower. We monitored reproductive phenology of 576 individual plants representing 305 species in 56 families in Sarawak, Malaysia. Observations continued for 53 mo from August 1992 and covered one episode of a general flowering cycle. Among 527 effective reproductive events during 43 mo, 57% were concentrated in the general flowering period (GFP) of 10 mo in 1996. We classified 257 species into flowering types based on timing and frequency of flowering. The most abundant type was “general flowering” (35%), which flowered only during GFP. The others were “supra-annual” (19%), “annual” (13%), and “sub-annual” (5%) types. General flowering type and temporal aggregation in reproductive events were commonly found among species in various categories of taxonomic groups, life forms, pollination systems, and fruit types. Possible causes for general flowering, such as promotion of pollination brought about by interspecific synchronization and paucity of climatic cues suitable for flowering trigger, are proposed, in addition to the predator satiation hypothesis of Janzen (1974) .     

Dynamics of island populations of Cypripedium calceolus in the Biebrza river valley (north-east Poland)

Three Cypripedium calceolus populations were observed in the Biebrza National Park (north-east Poland) for 11 years (1989–99). The populations differed in the number of genets and ramets and in their dynamics. Differences were in the proportion of flowering ramets, number of flowers, fruiting and recruitment. Fruiting and recruitment are low in populations of Cypripedium calceolus . Fruiting is probably pollinators limited, and recruitment is limited by environmental conditions. Changes in the size of populations of this species are caused by the following processes: changes in the number of genets (their appearance and death), changes in the size of clones (growth of individuals and their disintegration, death of individual ramets), dormancy of genets, and animal pressure. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 67–77.     

Interactive effects of introduced herbivores and post‐flowering die‐off of bamboos in Patagonian Nothofagus forests

Question: In November 2000, Chusquea culeou, a bamboo species dominating Andean forest understories in southern Argentina and Chile, massively flowered and died over a north‐south distance of ca. 120 km. Because bamboo is the major forage for large herbivores in these forests, we examined the interactive influences of the bamboo die‐off and herbivory by introduced cattle on understory and tree regeneration. Location: Lanín National Park, Argentina. Methods: Permanent plots, in and outside livestock exclosures, were installed in a Nothofagus dombeyi forest in patches of flowered and non‐flowered C. culeou. Plots were monitored over four years for changes in understory composition and tree seedling densities and heights. Results: After the C. culeou die‐off, new establishment of N. dombeyi was low, both with and without herbivory. Livestock alone directly increased N. dombeyi seedling mortality through physical damage. However, tree seedling browse ratings and height growth were interactively affected by bamboo flowering and herbivory; unfenced plots in flowered bamboo patches had the shortest seedlings, highest browse ratings, and lowest tree seedling annual growth rates. Understory cover was higher where livestock were excluded, and this effect was intensified in the patches of flowered bamboo. Neither herbivory nor bamboo flowering resulted in major changes in species composition, with the exception of Alstroemeria aurea. Conclusion: Effects of livestock on N. dombeyi regeneration were contingent on flowering of C. culeou. Prior to introduction of livestock, N. dombeyi regeneration was probably successful beneath canopy gaps during windows of opportunity following bamboo die‐off, but now livestock impede tree regeneration. Herbivory during bamboo withering periods also produces more open understories, particularly affecting palatable heliophyllous herb species such as Alstroemeria aurea. The results underscore the importance of assessing herbivore impacts on tree regeneration during relatively short periods of potential tree regeneration immediately following rare bamboo flowering and die‐off.     

Meta-analysis of phenotypic selection on flowering phenology suggests that early flowering plants are favoured

Flowering times of plants are important life-history components and it has previously been hypothesized that flowering phenologies may be currently subject to natural selection or be selectively neutral. In this study we reviewed the evidence for phenotypic selection acting on flowering phenology using ordinary and phylogenetic meta-analysis. Phenotypic selection exists when a phenotypic trait co-varies with fitness; therefore, we looked for studies reporting an association between two components of flowering phenology (flowering time or flowering synchrony) with fitness. Data sets comprising 87 and 18 plant species were then used to assess the incidence and strength of phenotypic selection on flowering time and flowering synchrony, respectively. The influence of dependence on pollinators, the duration of the reproductive event, latitude and plant longevity as moderators of selection were also explored. Our results suggest that selection favours early flowering plants, but the strength of selection is influenced by latitude, with selection being stronger in temperate environments. However, there is no consistent pattern of selection on flowering synchrony. Our study demonstrates that phenotypic selection on flowering time is consistent and relatively strong, in contrast to previous hypotheses of selective neutrality, and has implications for the evolution of temperate floras under global climate change.     

Selection on flowering time in a life‐cycle context

The main way in which plants can exert control over their local environment is by the timing of different events within their life cycles. Regarding timing of flowering as an integrated part of both the annual cycle and of the whole life cycle, rather than as an isolated event, has important implications for how we assess selection on timing of reproduction and interpret existing phenological patterns in perennial plants. I argue that: 1) we have little unequivocal evidence of pollinator‐mediated selection on flowering time, but perhaps more evidence of antagonist‐mediated selection; 2) much of selection on flowering time might occur before flowers have developed and after reproduction; 3) vital rates of non‐flowering individuals can influence the strength and direction of selection on flowering time, and 4) differences in the direction of selection on flowering date between years might well correspond to consistent selection on the mechanisms determining flowering time. Overall, a life cycle perspective on timing of flowering is likely to facilitate the identification of selective agents and the understanding of the complex mechanisms underlying spatial and temporal variation in selection as well as to enable more accurate predictions of responses to environmental change.