Flowering phenology in the central highland of Iceland and implications for climatic warming in the Arctic

The cool and short growing season that characterizes Arctic climates puts severe constraints on life cycles and reproduction in the Arctic flora. The timing of flowering is particularly critical and may affect both breeding system and reproductive success through the heavy penalties associated with later flowering. An 11-year study of 75 species in the central highland of Iceland showed that the onset of flowering varies greatly among years. The number of species in flower by the first week of July was closely correlated with air temperature (degree days above zero) in the preceding 5 weeks, but no correlations were found with degree days in May or with total degree days in the previous growing season. Time of snowmelt, which has widely been regarded as the environmental event initiating growth and flowering in alpine and arctic tundra, only had a significant effect when two exceptionally cold and late summers were included. The species studied, most of which have a wide distribution in the Arctic, are predicted to respond quickly to warmer spring and early summer temperatures. Accelerated phenologies may alter patterns of resource allocation, have implications for pollinators and pollinator-competition, and could increase the size, species richness and intraspecific genetic diversity of the soil seed bank. Received: 15 February 1997 / Accepted: 25 October 1997     

Flowering phenology of <Emphasis Type="Italic">Ulex europaeus</Emphasis>: ecological consequences of variation within and among populations

Reproductive phenology of gorse (Ulex europaeus L., Genisteae, Fabaceae) is unusual in that the onset and duration of flowering vary greatly among individuals within populations: some plants initiate flowering in autumn or winter and continue flowering through spring, others initiate flowering in early spring. To understand the origin of this diversity and its ecological consequences, we investigated flowering phenology of randomly sampled individuals from five different natural populations in Brittany (France). Reproductive success was evaluated for individuals with contrasting flowering patterns, from 16 natural populations. Flower production, pod production, seed production and seed predation were estimated. Plants initiating flowering in spring produced larger numbers of flowers and pods over a shorter period than plants flowering from winter to spring, which produced few flowers and pods at a time but over a longer period. Pod production of long-flowering plants did not differ significantly between winter and spring, but their pods were more intensively attacked by seed predators in spring than in winter. We discuss our results in relation to biotic and abiotic parameters. We postulate that long-flowering can be interpreted as a bet-hedging strategy, spreading the risk of pod failure (rotting or freezing) in winter and of seed predation in spring.     

Flowering plant phenology and weather in Alberta,Canada

Plant phenology can be used for biomonitoring climate change. The flowering of certain temperate zone plant species occurs in response to accumulated heat. Networks of observers presently provide data on the timing of the growth of native and crop plants to Agro-meteorological Departments in Europe and the United States. In Alberta, a phenological survey which began in 1987 records flowering times for 15 native plants, with about 200 volunteers contributing observations annually. Six years of data have been summarized and correlated with temperature measurements. The Alberta phenological data can provide a key to sound decision-making in two ways: by providing proxy data on key variables to which vegetation responds, and by providing a model for transforming simple weather data into biologically meaningful zones.     

Variability in nectar production and standing crop, and their relation to pollinator visits in a Mediterranean shrub

Nectar standing crops in flowers within an individual plant are often highly variable. This variability may be a by-product of the foraging activity of insect pollinators. Alternatively, plants may be selected to produce highly variable rewards to reduce consecutive visitation by risk-averse pollinators, thus diminishing within-plant pollen transfer. This study evaluated the roles of pollinator control vs. plant control over nectar variability in the bee-pollinated shrub Rosmarinus officinalis L. (Lamiaceae). We sampled nectar production, standing crop and pollinator visits in three shrubs of one population over 17 days during one blooming season. Nectar production rates were highly variable (CV = 1.48), and increased after rainy days. Nectar standing crops were even more variable (CV = 2.16), decreased with increasing temperatures, and increased with time since the last rain. Pollinator visit rates decreased with variability in nectar standing crops, increased with flower number per shrub, and were unaffected by variability in nectar production rates. Repeated sampling of marked flowers revealed no correlation between their nectar standing crops and production rates. These findings support the role of reward variance in reducing pollinator visits, but suggest that plants are not in complete control of this variability. Rather, plant-generated variability can be modified by intensive foraging activity of pollinators. Such pollinator control over nectar variability is likely to reduce the selective advantage of plant-generated reward variation. Handling Editor: Neal Williams.     

Infestation rate of argan fruit (Argania spinosa) by the Mediterranean fruit fly (Ceratitis capitata) in relation to phenology and maturation of the fruit

Argan (Argania spinosa (Sapotaceae)) forest covers about 700,000 ha in the Souss region (Morocco). The medfly Ceratitis capitata Wiedemann (Diptera: Tephritidae) is known to develop in argans. In this study, we examine variations in medfly infestation rates according to argan phenology from 1987 to 1990. Argan phenology varies greatly among trees, total fruit drop spreading from late March to mid-July; moreover, an exceptional fructification may occur in Autumn in some trees. From February to about May, the infestation rate is low except in fruit produced during autumnal fructification. In June and July, the rate increases greatly reaching 100% in some trees. Ripe and ripening argans are the most heavily infested; only 10% of immature fruits on the average are attacked by the medfly. Fallen argans are more heavily infested than fruit still on the tree. Productivity per argan averaged from one to three adult medflies. Our results demonstrate that argan forest in the Souss acts as an enormous reservoir of medflies for the nearby citrus orchards.

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Résumé L'arganeraie (Argania spinosa) couvre environ 700000 ha dans la région du Souss, au Maroc. La mouche méditerranéenne des fruits, Ceratitis capitata, est connue pour se développer dans les arganes. Des études ont été menées, de 1987 à 1990, pour étudier comment le taux d'infestation des arganes variait selon leur degré de maturation. La phénologie des arganiers varie fortement selon les arbres: par exemple la chute totale des fruits s'étale de fin mars à mi-juillet. De plus, une fructification peut être observée en automne mais elle ne concerne qu'un nombre limité d'arbres. De février à mai environ, le taux d'infestation des arganes reste faible, sauf pour les fruits produits pendant la fructification de l'automne précédent. En juin en juillet, le taux augmente fortement atteignant 100% dans certains arbres. Les arganes mûres ou mûrissantes sont les plus attaquées; seuls 10% en moyenne des fruits immatures sont infestés. Les fruits tombés sont toujours (sauf à la fin) nettement plus attaqués que ceux encore présents dans l'arganier. De une à trois Cératites adultes sont formées à partir d'une argane. Nos résultats démontrent que la forêt d'argane du Souss constitue un réservoir gigantesque de la Cératite pour les infestations des vergers d'agrumes proches.

     

Effects of parentage,prior fruit set and pollen load on fruit and seed production in Campanula americana L

Summary We conducted a controlled crossing experiment to examine the effects of maternal and paternal parentage, the size of the pollen load, and prior fruit production on the proportion of flowers that set fruit, seed number per fruit and seed weight in a natural population of Campanula americana. Effects due to the maternal parent were large for all measures of fruit and seed production, while the paternal parent had a significant effect only upon mean seed weight. As the number of prior fruits on the maternal plant increased the probability that a flower would produce a mature fruit, the number of seeds per fruit, and total seed weight per fruit all decreased. We found no effect of the size of the pollen loads used in this study on fruit or seed production. These results are consistent with those of other studies that suggest in natural plant populations maternal effects, especially environmental maternal effects, can have an overwhelming effect on fruit and seed production and on seed characteristics.     

The smaller fruit tortrix, Grapholita lobarzewskii: predicting the phenology of adult emergence

The smaller fruit tortrix, Grapholita lobarzewskii Nowicki, has become a major pest in the recent past in apple orchards north of the Alps. Nevertheless little is known about the biology and the behaviour of this species. The effect of temperature on post-diapause development and survival was studied under controlled conditions in order to establish a basis for the forecasting of adult emergence. Survival was found to be highest at 17 °C and lowest at 11.8 °C. The sex ratio did not differ significantly from 1:1. Based on a linear relationship between temperature and developmental rates, thermal thresholds of 9.6 °C and 9.8 °C were determined for females and males respectively. The thermal constants, i.e., the mean developmental times in physiological time units, were found to be 342 day-degrees for females and 317 day-degrees for males, suggesting a slight protandry. Based on the mean and the variance of the developmental times, a simple temperature-driven phenology model was built using a time-varying distributed delay. The model was validated by visually comparing the predictions with independent observations on adult emergence, and by calculating the temporal deviations of the predictions. In 4 out of 5 years the mean error was less than 3 days. The model was therefore found to give reliable forecasts of the emergence of G. lobarzewskii and can be used to determine the optimal time for the exposure of pheromone traps, the application of pheromone dispensers for mating disruption, and for the timing of insecticide applications.     

Aspects of floral structure and phenology in the genusConophytum (Mesembryanthemaceae)

The complex floral structure in the southern African genusConophytum (Mesembryanthemaceae; 77 spp.) is described in detail and assigned to three basic floral types, two of which can be divided into two subtypes. Correlations between structural features and phenological patterns are demonstrated and discussed in the family context as well as in relation to the systematic subdivision of the genus.     

Interspecific pollen transfer as a mechanism of competition: Consequences of foreign pollen contamination for seed set in the alpine wildflower,Polemonium viscosum

Summary Earlier studies showed that under natural conditions foreign pollen receipt is correlated with loss of seed set in populations of P. viscosum. Here, we report on experimental pollinations using foreign pollen from two co-flowering species that share pollinators with P. viscosum. Experiments were designed to ascertain whether improper pollen transfer causes reduced seed set under controlled conditions and, if so, to determine the stage at which interference occurs. In flowers pre-treated by pollination with either Mertensia or Castilleja pollen, adherence of subsequently deposited conspecific pollen grains was unaffected. However, conspecific grains had significantly lower germination and ovule fertilization success in flowers receiving foreign pollen than in controls. These results suggest that pollinator infidelity in high alpine plant communities can reduce female reproductive success of P. viscosum by inhibiting the performance of conspecific pollen grains. Negative effects of Mertensia and Castilleja pollen were comparable despite large interspecific differences in pollen grain morphology. Accordingly, susceptibility to foreign pollen interference appears to be an intrinsic property of P. viscosum that is not dependent on the unique structural properties of pollen produced by competitive species.     

A review of the life history,reproduction and phenology of Gracilaria

The basic life history of the red alga Gracilaria is of the three-phase Polysiphonia type but a number of species show deviations. Plants can bear both gametangia and tetrasporangia, either on separate parts of the thallus or on the same. Explanations include the in situ germination of tetraspores (allowing gametophytic thalli to be epiphytic on tetrasporophytes), the coalescence of spores or developing discs (resulting in chimaeras), mitotic recombination during cell division in the mature diploid thallus (resulting in patches of diploid male and female cells on the tetrasporophyte), a mutation eliminating the repression of female expression allowing haploid male plants to be bisexual and initial failure of cell walls to form during the development of tetraspores. Polyploids can be produced from plants with diploid gametangia. The sexes and phases are usually morphologically identical but gametophytes or their parts may be smaller. The growth rates of the sexes may differ and diploid juveniles may survive better than haploid. Neither polyploidy nor hybridization results in superior growth. The sex ratio is probably 1:1 but females may appear to be more abundant. Diploid and haploid phases are usually either about equal or diploids predominate, often depending on the type of substratum. At high latitudes reproduction peaks in late summer whereas in the tropics it may be high all year. In temperate regions growth rate is fastest and biomass highest in late summer; in the tropics peak biomass is mainly in the winter. Spermatia are effective for only a few h. Spores vary in size around 25 m, diploid ones usually being larger. Cystocarps or tetrasporangia in the field may not currently be releasing spores. In the laboratory spore release shows a diurnal rhythm, peaking during the night or day according to the species. All the above attributes are potentially important in planning and executing Gracilaria cultivation.     

Flower morphology, phenology and visitor patterns in an alpine community on Mt Olympos, Greece

Alpine vegetation, restricted to the top of high mountains, is among the vegetation types most endangered by global warming, currently predicted to raise temperatures from 1.1 to 6.4 °C, by the end of the century. Nevertheless, background information allowing evaluation of impacts is rather scarce for some geographic zones. Our study of an alpine community on the Plateau of Muses (2600–2750 m a.s.l.) of Mt Olympos, the highest mountain of Greece, conducted in 1993–1994, can provide such background information for the Mediterranean region. We studied features relating to phenology of flowering, floral morphology, distribution and abundance, and flower visitors of plant species that exhibit a biotic pollination syndrome. We identified dominant patterns and we further (i) explored the relative contribution of the plant features and abiotic factors studied in explaining the activity patterns of flower visitors, (ii) examined if flower and visitor traits of the alpine community match each other according to the classical pollination syndromes, and (iii) investigated whether the responses of individual plant species to the yearly climatic variability result into phenological patterns that characterize the whole community. The common strategy of the alpine community was for early flowering and long flower life span; consistently early flowering species were twice as many as late flowering ones, whereas floral longevity (estimated for 36 species) averaged 5.2 days. Duration of flowering (estimated for 57 species) averaged 18.2 days. Climatic variability affected onset of flowering; all late flowering species delayed their flowering during the year characterized by a humid and cold summer. Duration of flowering and floral longevity did not change in a consistent way. Hymenoptera (Aculeates) were the dominant flower visitors. They accounted for 43.3% of the visits recorded, with bumblebees making a little less than half. Diptera followed making 37.5% of the visits (most made by syrphid flies). There was a mismatch between flower-morphology and flower visitor traits; the alpine community had predominantly non-specialized, pale-colour flowers, which are traits assumed to correspond to Diptera dominance and absence of social bees. Visitation was influenced by flower abundance and duration of flowering; proportionately more Diptera, and proportionately less Hymenoptera visited species with short flowering periods and few flowers present in the field. In a number of cases, the phenological and flower visitor patterns of the community of Mt Olympos deviated from those observed in other alpine environments suggesting a mediterranean influence even at high altitudes.     

Sexual dimorphism and fruit production in a dioecious understory tree,Ilex opaca Ait.

Summary This study suggested that sexual selection is potentially an important factor in the maintenance of dioecy in the American holly, Ilex opaca (Aquifoliaceae). Sexual dimorphisms in flower production and phenology were highly significant in this understory tree. On average, individual males produced 7.4 times as many flowers as did female trees. Staminate flowers lasted only a single day, whereas pistillate flowers lasted 3–4 days, during which they showed no significant decline in their ability to produce fruit after pollination. Individual male trees opened their flower buds asynchronously during the season, maximizing the number of days they were in flower. Individual females opened their buds more synchronously, maximizing their floral display at one point in time. Females produced fruits in numbers that were somewhat less than proportional to their flower production. Fruit development was initiated from only 38.9% and 69.5% of pistillate flowers in 1987 and 1988, respectively. By the time of ripening, an average female had lost 62.3%, 24.3%, and 11.1% of its initial fruit crop in 1986, 1987, and 1988, respectively. The proportion of fruit lost in 1986 was independent of the number of fruit that initially began development. In 1988, artifically supplementing pollen to a large number of flowers failed to increase either fruit or seed production relative to control branches with unsupplemented flowers. This suggested that resource levels were likely more important than pollen availability in limiting female reproductive success. These observations on I. opaca were consistent with the expectations for a population in which male reproductive success continues to benefit from continued pollinator service and female reproductive success does not.     

Flowering phenology, display size, and fruit set in an understory dioecious shrub, Aucuba japonica (Cornaceae)

I investigated the effects of display size and flowering phenology on fruit set in Aucuba japonica, an understory dioecious shrub pollinated by opportunistic insects. Natural variations in display size, flowering phenology, and fruit set were monitored in 1997. A hand-pollination experiment was also conducted to check whether pollen limitation was a factor in fruit set in the field. Increases in floral display size did not affect fruit set; the proportion of flowers that set fruit was almost constant irrespective of the total number of flowers per inflorescence, the total number of inflorescences per plant, and the total number of flowers per plant. The hand-pollination experiment showed that fruit set was not pollen limited despite the low mating probability that resulted from the combination of dioecism and the species' dependence on opportunistic pollinators. This was due, in part, to the fact that female flowers did not have a predetermined period of receptivity, but instead remained receptive until they received pollen. In contrast, flowering phenology did affect fruit set. Fruit set was most abundant when male and female flowering was most abundant. This suggests there was some degree of pollen limitation during the part of the flowering season when male flowers were scarce.     

Density effects of flowering phenology and mating potential in Nicotiana alata

Summary We investigated effects of plant density on floral phenology and potential mating in artificial populations of the outcrossing ornamental Nicotiana alata planted at three densities. Path analysis revealed that increasing plant density yielded significantly earlier peak flowering dates, significantly earlier last flowering dates, and significantly lower plant biomass. Direct effects of density on final flower number were not significant. Variation among replicate plots for first date of flowering was larger than variation among densities, indicating that factors other than density influence floral initiation.We did not record actual mating, but determined from phenological data the number and identity of potential mates. Increased density had several effects on potential mating patterns and on potential N_e, effective population number. At high density, fewer focal plants flowered for shorter durations. This led to less overlap in flowering time among plants, decreasing the number of potential parental combinations possible among the progeny. Two outcomes of high density, the lower total number of plants flowering and the lower number of plants flowering at most census dates, tended to reduce potential N_e. In contrast, it was low density, where variance in flower number was greatest, that was most likely to yield the greatest reduction in N_e due to variance in progeny number.At high density the potential for assortative mating among tall plants was much greater and occurred later than among large plants at low density. Much of the potential high density assortative mating occurred late in the phenology of individual plants, when there was likely to be lower fruit set.We discuss how ecological agents that alter flowering phenology can potentially alter the genetics of populations, the level and timing of assortative mating and, if genetic variation for response to such ecological agents exists, the potential selection regime.     

Population response of the twospotted spider mite,Tetranychus urticae, to host phenology in corn and peanut

In northeastern North Carolina, outbreaks ofTetranychus urticae Koch on commercial corn and peanut plantings were observed to coincide with flowering and fruiting of the crop host. In greenhouse studies, when equal mite numbers were started on plants in either vegetative or reproductive growth stages, populations increased significantly more after 3–4 weeks on reproductive plants of both corn and peanut. This direct response of mite populations to differences in plant phenology appears to be an important component in the population dynamics ofT. urticae. The importance of this effect in understanding mite outbreaks on corn and peanut is discussed, especially in reference to the corn-peanut agroecosystem in North Carolina.

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Réponse de populations deTetranychus urticae Koch., aux phénologies du maïs et de l'arachide

Résumé Les populations de tétraniques sont souvent associées à la floraison et à la fructification des plantes attaquées. Dans les agrosystèmes maïsarachide du N.E. de la Caroline du Nord, l'augmentation rapide de populations deT. urticae a été observée lors de la maturation des épis mâles du maïs et du maximum de floraison de l'arachide. Bien que ces observations suggèrent une relation causale avec la phénologie des plantes hôtes, il est difficile de déterminer dans les conditions de la nature si l'augmentation des populations d'acariens sur les stades reproducteurs des plantes est due à une réponse à la phénologie des cultures ou à une autre cause, comme le passé de la population ou une immigration. Des expériences ont été réalisées dans une serre pour mettre en évidence l'action de la phénologie du maïs et de l'arachide sur les populations d'acariens en maîtrisant ces autres hypothétiques facteurs. Des effectifs identiques d'acariens femelles ont été libérés sur des plantes à des stades tant végétatifs que reproductifs, et ensuite laissés sans interventions pendant plusieurs semaines, au bout desquelles les effectifs sur chaque plante ont été dénombrés. Sur les deux hôtes, les populations sur plante à un stade reproducteur avaient augmenté significativement plus que sur les plantes de même espèce à un stade végétatif. Ces résultats montrent queT. urticae répond directement à des différences entre plantes à un stade végétatif et à un stade reproductif. Cette réponse semble jouer un rôle important dans la dynamique des populations deT. urticae.

     

Ecological aspects of Sargassum muticum (Fucales,Phaeophyta) in Baja California,Mexico: reproductive phenology and epiphytes

The reproductive phenology and epiphytic macroalgae of Sargassum muticum were studied through an annual cycle (September 1987 to November 1988) at two sites on the northwestern coast of Baja California, Mexico, which were subjected to different degrees of wave exposure. Sargassum muticum is a brown alga of Japanese origin, now considered a permanent member of the marine flora of Baja California. A similar reproductive development was observed at both sites, with a maximum percentage of reproductive plants from May to July (spring–summer) and minimum from December to March (winter). Reproductive plants were found throughout the year. A total of 48 species of epiphytes were identified and seasonal variation in their diversity was observed. The greatest diversity was found at the more protected site.     

Effect of in vitro and in vivo colchicine treatments on pollen production and fruit set of melon plants obtained by pollination with irradiated pollen

Haploid/doubled haploid (DH) technology can aid plant breeding programs by accelerating production of homozygous lines, provided enough viable DH progeny can be obtained from diverse haploid genotypes. In cases where there is a low frequency of spontaneous doubling, chromosome doubling procedures are required to achieve fecundity. We produced 63 parthenogenetic melon plantlets via pollination with γ-irradiated pollen, cloned them by nodal cuttings, and tested the effects of in vitro and in vivo colchicine treatment on survival, ploidy, pollen production, and fruit recovery. The most effective procedure was in vitro exposure of 3 cm shoot tip explants to 500 mg/l colchicine for 3 h. This treatment gave 83% survival of explants and 26% conversion to diploidy. Fruit recovery rate was 60% among plants with good pollen production. In vivo exposure of the tops of young plants to 5000 mg/l for 2 and 4 h yielded some fruits but also resulted in less survival and more morphological abnormalities. Strategies for recovery of progeny from parthenogenetic melon plants are recommended. To our knowledge, this study represents the first comprehensive study of recovery of fruits and viable seeds from parthenogenetic melon plants.     

Floristic composition and floral phenology of the Mediterranean batha of Ariel,Samaria

Field observations over a three-year period were used to document the floristic composition, flowering and fruiting sequences of the Sarcopoterium spinosa Mediterranean batha of Ariel. This plant community was found to be species-rich, with high percentages of annuals, bushes and geophytes, low percentages of herbaceous plants, climbers and trees, and ten species considered rare in Israel. Typical Mediterranean characteristics were observed in the S. spinosa batha: spring flowering reaching a peak in April, small between-year variation in flowering and fruiting, most species having Mediterranean distribution, and partial resistance to fire.     

The phenology ofProcladius crassinervis (Zett),Procladius signatus (Zett) andProcladius choreus (Mg) (Diptera,Chironomidae) at Lake Hald,Denmark

In an investigation of the phenology of the chironomid species of Lake Hald, Denmark, a very late flight activity (September–November) of the univoltine speciesProcladius choreus (Mg.) was observed. Among observations of the phenological patterns of univoltine chironomids inhabiting lakes of the Baltic type in Denmark, the phenological pattern ofProcladius choreus stands alone (JÓNSSON, 1987; LARSEN, 1991). Further two species of the genusProcladius Skuse,viz. Procladius crassinervis (Zett) andProcladius signatus (Zett), were found in the investigation and the phenological pattern of the three congeneric species was clearly allochronic, withProcladius crassinervis flying in spring,Procladius signatus flying in summer andProcladius choreus flying in autumn.