开花时间与伴生种对鹤首马先蒿传粉和生殖成功的影响

开花物候是物种间相互作用的重要生活史特征和适合度因子,在全球气候变化的背景下而备受关注.为探讨开花时间如何存种内和种间水平上影响植物的传粉和生殖成功,我们连续3年(2003-2005)对不同花期和伴生种存在情况下的鹤首马先蒿(Pedicularis gruina)的传粉者访花忠实度、受粉率、坐果率、单果种子产量和果实被啃食频率进行了比较研究.结果表明鹤首马先蒿的坐果率主要受其传粉环境的影响:在没有伴生种时,不同时期鹤首马先蒿的坐果率没有显著差异,34-38％的花可以坐果;在有伴生种存在时既可以显著提高其坐果率,也可显著降低其坐果率,这取决于传粉者类型以及伴生种密穗马先蒿(P.densispica)花期的差异.密穗马先蒿具有花蜜和花粉双重报酬,在群落中可以作为主体物种吸引传粉者并间接促进与其伴生的鹤首马先蒿的传粉和生殖成功.同样无蜜的管花马先蒿(P.siphonantha)和鹤首马先蒿伴生,则是通过提高群落水平对传粉者的吸引力进而直接促进鹤首马先蒿的传粉和生殖成功.此外,研究结果也表明开花时间对坐果率没有显著影响,但是显著影响单果种子产量和果实被啃食的频率.在相同的传粉条件下早花期植株单果种子产量显著高于晚花期的种子产量,同时早花期的果实被啃食的频率显著增加.     

SEED SET AND SEED PREDATION IN PEDICULARIS FURBISHIAE,A RARE ENDEMIC OF THE ST. JOHN RIVER,MAINE

We measured seed set and losses due to mammal and insect predation during 1983 in seven northern Maine populations of Furbish's Lousewort (Pedicularis furbishiae S. Wats.; Scrophulariaceae), a perennial herb endemic to the banks of the St. John River. Only about a quarter of initiated flowers developed a capsule with a full complement (mean = 25) of seeds. Secondary (lateral) inflorescences matured a smaller proportion of capsules than primary (terminal) inflorescences. About 58% of the initiated flower primordia were aborted or did not develop mature seeds by early September; this was probably not due to pollinator limitation. Several agents could have reduced seed production. Although spittlebugs infested between 3 and 30% (average 20%) of developing flower scapes by population, they did not depress the number of inflorescences initiated or the number of capsules matured. In contrast, 74% of capsules containing a lepidopteran seed predator (the plume moth Amblyptilia pica [Walsingham], newly discovered as a seed predator on P. furbishiae) had no seeds (mean = 3 seeds). Seed predation affected 39% of the potentially maturable capsules. In 6–30% of all capsules preyed upon, the lepidopteran was parasitized by an ichneumonoid wasp larva; these capsules matured about 7 seeds. Herbivory by rabbits, rodents, and deer also reduced seed set, eliminating 98% of the scapes at one site, and averaging 40% for other populations. Mammals and insects together removed at least 62% of maturable seeds. Populations varied widely in predation and seed set. Predispersal seed loss in P. furbishiae, coupled with slow seedling growth and occasional catastrophic mortality from ice scour and erosion, could limit population growth in this rare endemic.     

Taxonomic position and distribution of Atriplex lapponica (Chenopodiaceae)

The arctic, halophytic species Atriplex lapponica Pojark has flowers of two kinds: perfect ones surrounded by a pentamerous perianth and pistillate flowers enclosed in bracteoles. Thus, A. lapponica is closely related to the species of sect. Dichospermum Dumort, not to species of sect. Teutliopsis Dumort. as generally believed.     

Reproductive Ecology of Understory Species in a Tropical Montane Forest in Costa Rica1

The reproductive ecology of nine hermaphroditic understory species in a tropical montane Quercus forest was studied at two sites (2300 and 2600 m elev.) in the Cordillera de Talamanca, Costa Rica. Flower life span, studied in six species, averaged 4.4 d. This is longer than flower life spans found in the Monteverde cloud forest (2.7 d) and comparable to flower life spans found for arctic and alpine species. We studied the breeding system in five species and found no self-incompatible species. Four species proved self-compatible, and three of these showed autogamy. The main diurnal insect pollinator was the bumblebee Bombus ephippiatus. Natural fruit set was low (8-32%) in six species with few seeds per fruit, while two many-seeded species showed a high rate of fruit set (90 and 96%). The incidence of pre–dispersal seed predation was high; the percentage of seeds infested in four species ranged from 8 to 56 percent.     

Resource limitation and pollen source (self and outcross) affecting seed production in two louseworts, Pedicularis siphonantha and P. longiflora (Orobanchaceae)

Two animal-pollinated hermaphrodite plants, Pedicularis siphonantha and P. longiflora , have been used to investigate factors limiting seed production in natural populations. To evaluate the potential seed abortion due to resources limitation, seed development has been observed and seed count conducted twice. Seed production per capsule has been compared when flowers have been removed and in a control group. Open pollination has been investigated and pollen supplementation undertaken to estimate the possibility of pollen limitation. Results show that seed abortion is frequent. Stigmatic pollen load is significantly higher than ovule number per ovary under open pollination for both species. Additional self and outcross pollen did not affect seed production. Flower removal significantly increases seed production per capsule, which indicates that seed production of the studied species is limited by available resources. To detect differences in seed production between flowers pollinated by self and outcross pollen, hand pollination of bagged flowers has also been conducted in natural populations of the two Pedicularis species. Compared with open pollination, hand-pollinating self-pollen decreases, while outcross pollen increases seed production per capsule. Such results suggest that inbreeding depression in the two self-compatible species may also result in partial seed abortion under open pollination if mixed pollen is deposited on the stigma. Our results also suggest that pollen interference plays an important role in low female fertility in the two Pedicularis species.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 83–89.     

The comparison of selected life-history characteristics of Mesembryanthema species occurring on and off Mima-like mounds (heuweltjies) in semi arid southern Africa

Selected life-history characteristics of Mesembryanthema species occurring on and off mima-like mounds (heuweltjies) were compared at Tierberg in the southern Succulent Karoo, South Africa. Seed production, the number of seeds retained in seed capsules and germination behaviour were measured in fourteen species of Mesembryanthema which were representatives of two sub-families: the Ruschioideae and the Mesembryanthemoideae. Species occurring onheuweltjies had low overall germination and high seed retention compared with those species occurring offheuweltjies. A strong negative correlation was found between the degree of seed retention and maximum germination. It was proposed that rapid germination and low seed dormancy in species occurring on the flats would be a viable strategy where competition for safe recruitment sites is strong. Species occurring on the flats are long-lived woody shrubs, thus the resulting high levels of seedling mortality probably do not have a major impact on population dynamics.Heuweltjies are sites of intense disturbance. Plant species that occur onheuweltjies can also occur offheuweltjies in disturbed vegetation. Opportunistic life-history characteristics allow these species to colonise disturbed areas.Heuweltjie species thus act as pioneers in a successional process which results ultimately in the re-colonisation of the long-lived flats species. Phylogenetic factors may also determine the species distributions. Members of the Mesembryanthemoideae did not occur offheuweltjies in undisturbed areas. Most Ruschioideae were found on the flats but the few species that occurred onheuweltjies shared similar characteristics with the Mesembryanthemoideae. The primitive life-history characteristics exhibited by the Mesembryanthemoideae enable them to exploit disturbed conditions.     

Water relations, temperatures, and mineral nutrients in Pedicularis dasyantha (Scrophulariaceae) from Svalbard, Norway

The arctic vascular hemiparasite Pedicularis dasyantha has a lower diffusion resistance for water vapour than any associated species. Excised leaves die within one hour of drought stress under laboratory conditions, in spite of a high succulence. Transpiration rates of P. dasyantha in the field are almost two times higher than in Dryas octopetala , which is the most important host within investigated sites.
Diffusion resistances are higher in inflorescences than in leaves, probably because the dense pubescence reduces transpiration rates and raises temperatures in inflorescences to 14°C above air temperature on clear days.
Pedicularis dasyantha contains significantly more N, P, and especially K than its host. The high rate of transpiration probably enables the hemiparasite to capture a larger fraction of nutrients supplied by the xylem of its host.     

Flower herbivory and seed predation in Silene vulgaris (Caryophyllaceae): effects of pollination and phenology

Pollination and seed predation were studied in Silene vulgaris populations during two seasons, one with much lower pollinator abundance than the other. Among the pollinators, noctuid moths of the genus Hadena also acted as seed predators. Nectar-foraging female moths oviposited in flowers, and their larvae consumed flowers and seed capsules.
Despite a lower percentage of pollinated flowers in the year of low pollinator abundance, similar numbers of flowers set fruit in both years, because fewer flower buds and flowers were eaten by Hadena larvae during the year of low pollinator visitation. The number of seed capsules preyed upon was also lower in the year with low pollinator abundance, resulting in a higher seed set. The positive correlation between the percentage of pollinated flowers and the percentage of seed capsules destroyed was also observed when comparing flowers opening in different parts of the season.
Early flowering plant individuals had the same pollination success but suffered higher seed predation than late flowering ones. Selection for maximized pollination success through synchronous flowering, is probably the main reason for the compressed flowering period in 5. vulgaris , but the high level of predation early in the season may further increase the reproductive success of synchronous flowering individuals.     

The biology and life history of arctic populations of the littoral mite Ameronothrus lineatus (Acari, Oribatida)

The present study attempts to elucidate possible microevolutionary adaptations of life-history traits of high-latitude populations of the holarctic, littoral oribatid mite Ameronothrus lineatus by comparing arctic and temperate populations. Additionally, the paper provides an overview of the limited research on general ecology and population biology of arctic populations. In the Arctic the larviparous A. lineatus has a 5-year life cycle (larva-to-larva), and adults survive a further 2–3 years. High survival to maturity is consistent with a low lifetime reproductive output of ca. 20 larvae. The life history can be regarded as an extreme version of the typical oribatid life history. However, several life-history features suggest specific adaptations of arctic populations. In particular, the pre-moult resting stage is synchronized with the warmest part of the arctic summer, which shortens this vulnerable part of development. High reproductive investment by females at relatively low temperatures may represent a physiological adaptation to the cool arctic summer. Finally, prolonged cold exposure positively affects reproduction and survival the following summer, suggesting adaptation of the species to the highly seasonal arctic environment. On the other hand, the ability of all life-cycle stages to overwinter, and a flexible life history with the species being able to take advantage of favourable climatic conditions to accelerate development and larviposition, seem to be ancestral features. Thus, the success of A. lineatus in arctic habitats is probably attributable to a combination of derived and ancestral life-history traits. Studies of conspecific temperate populations are required to elucidate further local adaptations of arctic populations.     

The biology and life history of arctic populations of the littoral mite Ameronothrus lineatus (Acari, Oribatida)

The present study attempts to elucidate possible microevolutionary adaptations of life-history traits of high-latitude populations of the holarctic, littoral oribatid mite Ameronothrus lineatus by comparing arctic and temperate populations. Additionally, the paper provides an overview of the limited research on general ecology and population biology of arctic populations. In the Arctic the larviparous A. lineatus has a 5-year life cycle (larva-to-larva), and adults survive a further 2-3 years. High survival to maturity is consistent with a low lifetime reproductive output of ca. 20 larvae. The life history can be regarded as an extreme version of the typical oribatid life history. However, several life-history features suggest specific adaptations of arctic populations. In particular, the pre-moult resting stage is synchronized with the warmest part of the arctic summer, which shortens this vulnerable part of development. High reproductive investment by females at relatively low temperatures may represent a physiological adaptation to the cool arctic summer. Finally, prolonged cold exposure positively affects reproduction and survival the following summer, suggesting adaptation of the species to the highly seasonal arctic environment. On the other hand, the ability of all life-cycle stages to overwinter, and a flexible life history with the species being able to take advantage of favourable climatic conditions to accelerate development and larviposition, seem to be ancestral features. Thus, the success of A. lineatus in arctic habitats is probably attributable to a combination of derived and ancestral life-history traits. Studies of conspecific temperate populations are required to elucidate further local adaptations of arctic populations.     

Evidence of a component Allee effect driven by predispersal seed predation in a plant (Pedicularis rex,Orobanchaceae)

A small or sparse population may suffer a reduction in fitness owing to Allee effects. Here, we explored effects of plant density on pollination, reproduction and predation in the alpine herb Pedicularis rex over two years. We did not detect a significant difference in the pollination rate or fecundity (fruit set and the initial seed set) before predation between sparse and dense patches in either year, indicating no pollination-driven Allee effect. However, dense patches experienced significantly fewer attacks by predispersal seed predators in both years, resulting in a significantly decreased realized fecundity (final seed set), suggesting a component Allee effect driven by predispersal seed predation. Predation-driven Allee effects have been predicted by many models and demonstrated for a range of animals, but there is scant evidence for such effects in plants. Our study provides strong evidence of a component Allee effect driven by predation in a plant species.     

SPATIAL PATTERNS OF SEED DISPERSAL AND POSTDISPERSAL SEED PREDATION OF CRYPTANTHA FLAVA (BORAGINACEAE)

I examined the spatial patterns of seed dispersal and postdispersal seed predation of the semidesert perennial Cryptantha flava (A. Nels.) Payson (Boraginaceae) at two sites in north-eastern Utah. Most flowers mature only one seed (nutlet) which is permanently retained within a pubescent calyx. The calyx and enclosed seed abscise from the plant as a unit. These dispersal units are effectively dispersed by wind as evidenced by the highly directional seed shadows and the long distances some of them travel (up to 31.3 m). Potential seed predators at the sites include five species of rodents, of which Peromyscus maniculatus is the most common, and two species of ants, Pogonomyrmex occidentalis and an undescribed species of Conomyrma. There were no strong spatial patterns of postdispersal seed predation. More seeds were removed from dishes placed at the bases of fruiting adults than from dishes ≥ 1.0 m away in one of three experiments. More seeds were removed from under shrubs or clumps of grass than in the open in one of four experiments. After 3–4 days, there was a consistent tendency for more seed removal from high density (75 seeds per .25 m²) quadrats than from low density (75 seeds per 6.25 m²) quadrats, but the difference was not always significant. There was a similar nonsignificant difference between high- and low-density quadrats exposed for 21 days. The pubescent calyx greatly discourages seed predation by ants, and probably also reduces predation by rodents. In addition, by increasing the surface area of the dispersal unit, the calyx may facilitate dispersal by wind.     

Seed germination of high mountain Mediterranean species: altitudinal,interpopulation and interannual variability

The germination response of 20 species from high altitude Mediterranean climates, most of them rare endemics, was studied. Our main goal was to model the germination response of a complete set of Iberian high mountain species. The effect of temperature and other parameters, such as spatial and temporal short gradients, on germination were also evaluated. Some seed features (mass and size) were also related to the germination response. Finally, we tested the effect of cold-wet stratification pretreatment when germination was low under natural conditions. Seeds were collected at four locations from 1,900 to 2,400 m a.s.l. in the Sierra de Guadarrama (Spanish Central Range) over two consecutive growing seasons (2001–2002) and submitted to different temperatures and a constant photoperiod of 16 h light/8 h darkness. Most plants readily germinate without treatment, reaching an optimum at relatively high temperatures in contrast to lowland Mediterranean species. Seeds seem to be physiologically prepared for rapid germination even though these plants usually face very intense summer droughts after ripening and dispersal. Germination was also highly variable among altitudes, populations and years, but results were inconsistent among species. Such flexibility could be interpreted as an efficient survival strategy for species growing under unpredictable environments, such as the Mediterranean climate. Finally cold-wet stratification increased germination capacity in five of nine dormant species, as widely reported for many arctic, boreal and alpine species. In conclusion, high mountain Mediterranean species do not differ from alpine species except that a relatively high number of species are ready to germinate without any treatment.     

Revisions of Pterogastra and Schwackaea (Melastomataceae: Melastomeae)

The closely related neotropical melastomean genera Pterogastra and Schwackaea are revised, the species illustrated, and their distribution mapped. Schwackaea is maintained as a monotypic genus because it is readily distinguished from all other melastomes by its 8-winged capsule. This weedy little plant occurs from Mexico throughout Central America, chiefly on the Pacific slope, to northern Colombia and on Cocos Island and grows in natural or man-made savannas from sealevel to 2000 m altitude. Two species are recognized in Pterogastra and one new combination, P. divaricata spp. glabra , is made. Pterogastra is characterized by distinctly 4- or 5-winged capsules, unlike those of Schwackaea or any other New World melastome. The more widespread species occurs in Colombia, Venezuela, Guyana, Ecuador, and Peru at altitudes between sealevel and 2600 m in natural or disturbed grasslands. The second species is endemic in southwestern Venezuela and is confined to natural savannas at low altitudes.     

The predation of predispersed Juncus squarrosus seeds by Coleophora alticolella (Lepidoptera) larvae over a range of altitudes in northern England

Seed production by the rush Juncus squarrosus and its subsequent predation by the larvae of the moth Coleophora alticolella were measured at a series of sites at different altitudes in northern England. The density of seeds produced is progressively reduced with increasing altitude. The density of larvae was greatest in the middle region of the transect. Seed predation was therefore highest at about 300 m. Predation was reduced to nil above 520 m, where the larvae were absent, but remained between 40 and 50% below 250 m. The density of predispersed seed surviving was greatest at the lowest site where the highest density of seeds was produced and predation was moderate. The lowest densities of surviving seeds were found around 300 m and also above 600 m where there was no predation but few seeds were produced. A similar density survived around 520 m as at 245 m, although the density of seed produced at 520 m was less than 30% of that at 245 m. The results are discussed in relation to the population dynamics of the moth in different parts of its altitudinal distribution.     

Adult body size = <Emphasis Type="Italic">f</Emphasis> (initial size + growth rate × age): explaining the proximate cause of Bergman’s cline in a toad along altitudinal gradients

Animals that exhibit indeterminate growth obey such a functional relationship: adult body size = f (initial size + growth rate × age). Using this framework, we investigated how and why body sizes of a toad species (Bufo andrewsi) covaried across six altitudes (760–2,100 m) in western China. Towards high altitudes, toads tended to produce large eggs, attain large sizes at metamorphism and have great average age, but grow slowly. This indicated that the former three variables contributed more to the observed altitudinal increase in body size than did the last one. The altitudinal variation in these life-history traits should be adaptive to increased climate harshness and decreased predation risks at higher altitudes. We suggest that the relative significance of responses of these size-related parameters to local environments may provide critical cues to explaining considerable variability in geographic size pattern among ectothermic vertebrates.     

Divergence among arctic and alpine populations of the annual, Koenigia islandica: morphology, life-history, and phenology

Arctic and alpine habitats occur along complex environmental gradients, and over an extensive geographical range. Despite some selective forces common to these habitats, evolutionary divergence among populations of arctic and alpine plants along this gradient is expected. Of particular significance, both in the context of life-history theory and for implications of climate change, are the few annual species that have adapted to the constraints of an unpredictable, short growing season. In this study, morphological, life-history and phenological characters were found to differ significantly among six widely distributed populations of the arctic-alpine annual Koenigia islandica. On the basis of morphology and life-history traits, populations from high latitudes, with the exception of Svalbard, performed better in simulated arctic conditions, whereas the low latitude alpine plants from Colorado showed enhanced performance under simulated alpine conditions. On the basis of phenology, the six populations can be clearly grouped into arctic, high latitude alpine and alpine populations: arctic plants were found to develop and flower earliest; alpine plants latest. Because these results were obtained using seeds harvested from plants first grown through a complete generation in growth chambers, they indicate strong genetic differentiation. We discuss possible adaptive explanations for observed differences among the six geographically divergent populations.     

LIFE-HISTORY EVOLUTION IN GUPPIES (POECILIA RETICULATA) 6. DIFFERENTIAL MORTALITY AS A MECHANISM FOR NATURAL SELECTION

We have previously reported a correlation between the life-history patterns of guppies and the types of predators with which they coexist. Guppies from localities with an abundance of large predators (high predation localities) mature at an earlier age and devote more resources to reproduction than those found in localities with only a single, small species of predator (low predation localities). We also found that when guppies were introduced from a high to low predation locality, the guppy life history evolved to resemble what was normally found in this low predation locality. The presumed mechanism of natural selection is differences among localities in age/size-specific mortality (the age/size-specific mortality hypothesis); in high predation localities we assumed that guppies experienced high adult mortality rates while in the low predation localities we assumed that guppies experienced high juvenile mortality rates. These assumptions were based on stomach content analyses of wild-caught predators and on laboratory experiments. Here, we evaluate these assumptions by directly estimating the mortality rates of guppies in natural populations. We found that guppies from high predation localities experience significantly higher mortality rates than their counterparts from low predation localities, but that these higher mortality rates are uniformly distributed across all size classes, rather than being concentrated in the larger size classes. This result appears to contradict the predictions of the age/size-specific predation hypothesis. However, we argue, using additional data on growth rates and the probabilities of survival to maturity in each type of locality, that the age-specific mortality hypothesis remains plausible. This is because the probability of survival to first reproduction is very similar in each type of locality, but the guppies from high predation localities have a much lower probability of survival per unit time after maturity. We also argue for the plausibility of two other mechanisms of natural selection. These results thus reveal mortality patterns that provide a potential cause of natural selection, but expand, rather than narrow, the number of possible mechanisms responsible for life-history evolution in guppies.     

Predation and life-history variation in Poecilia reliculata (Cyprinodontiformes: Poeciliidae)

Synopsis Reznick and Endler investigated natural variation in life-history traits of populations of Trinidad guppies exposed to one of three intensities of predation: (i) high predation directed primarily at adults, (ii) moderate predation directed primarily at juveniles, and (iii) low predation. They were able to document significant interpopulational differences in life-history traits associated with this differential predation on a trait-by-trait basis. However, the present extended multivariate analysis indicates that (1) life-history traits do not differ significantly between populations exposed to moderate versus low predation, although both differ greatly from high-predation populations; (2) life-history variation is strongly unifactorial; and (3) despite the importance of predation effects, approximately 17% of the variation in life-history variables cannot be accounted for by predation intensity. Residual variation has no obvious geographical patterns, but instead seems to reflect local environmental variability. Life-history differences between predation regimes are consistent with residual patterns of variation within regimes, suggesting that local variation provides the raw material for extrapolation in response to predation, but also that it influences the direction of correlated change in life-history traits.