Seed germination ecology of Polymnia canadensis (Asteraceae), a monocarpic species of the North American Temperate Deciduous Forest

The seed germination of Polymnia canadensis was studied in relation to its wide variation in life history and its ecology as a mostly facultative biennial, a life cycle type that is rare in stable forest habitats in eastern North America. The seed dormancy and germination characteristics of P. canadensis were like some other herbaceous woodland species of diverse life cycle types. That is, its seeds have physiological dormancy, and they (1) germinated to higher percentages after cold stratification, (2) germinated better in light than in darkness and (3) formed a persistent soil seed bank. Since its seed dormancy and germination characteristics are not uncommon and are not particularly associated with any one life cycle type in woodlands, those properties do not contribute to the rarity of its life cycle type in deciduous forests. These properties are also found in many facultative biennials of open, disturbed habitats and in other Asteraceae of various life cycle types. In addition, the germination phenology of P. canadensis differed between mesic and dry field sites in central Kentucky. At the dry sites, most of the yearly germination occurred in fall, while at the mesic sites most germination occurred in the spring due to being inhibited during fall. A reciprocal seed transplant experiment conducted in two different years demonstrated that the cause of this difference was environmental and that there was no source x site interaction; i.e., no difference between sources in phenotypic plasticity for germination phenology. Absence of genetic effects was also shown by no consistent pattern in the few differences between seed sources in seed incubation laboratory experiments and in a germination phenology experiment in glasshouse flats. Exposure of seeds to a high ratio of far-red/red light prior to or during incubation did not inhibit germination. Neither did changes in leaf litter thickness qualitatively alter germination phenology in field quadrats or in glasshouse flats. Thus, germination in fall at mesic sites was inhibited by other unknown factor(s), perhaps interacting with the effects of closed tree canopy and/or thick leaf litter.     

Phenology and common garden and reciprocal transplant studies of Polymnia canadensis (Asteraceae), a monocarpic species of the North American Temperate Deciduous Forest

Polymnia canadensis, an herbaceous plant species ofthe North American Temperate Deciduous Forest heretofore reported as beingperennial, is shown to be primarily monocarpic. In common garden experiments,the majority of plants grown from seeds or seedlings collected in Kentucky,Alabama, Virginia, Ohio, and Missouri were monocarpic, but some were dicarpic,tricarpic or polycarpic. Observations in natural populations of P.canadensis suggested that there is variation in the life history ofthis species throughout its geographic range. Anthesis date and height ofplantsat anthesis were significantly different between mesic and dry field sites incentral Kentucky populations, but common garden and reciprocal transplantexperiments showed that these were due mainly to environmental (and not togenetic) effects. The latter was also true for age of maturity and rate ofchange in plant height. Lack of significant site × source interaction forthese characters indicated that plants did not differ in their plasticresponsesbetween sources at each site. Winter annual, biennial, triennial, and dicarpicand tricarpic perennial life histories occurred within populations in bothcommon garden and reciprocal transplant experiments. In the latter experiments,most matured plants were either biennials (most common) or winter annuals(second most common), with the order reversed in the former experiments due toamore benign environment favoring earlier maturity. The extensive phenotypicplasticity displayed by P. canadensis for phenology andlife history enables the species to inhabit a wide range of environments.     

The demography of the annual halophyte Spergularia marina on a baltic seashore meadow

The survivorship and reproduction of the annual halophyte Spergularia marina was studied during four years on three different sites on a Baltic seashore meadow. The survivorship showed variation both between years and between sites. The mortality rate was not density-dependent in spite of very high densities in some years. At two of the sites the seed output was related to density, but individuals differed in their reproductive capacity. Limitation of the flower production was the main reason for this difference. At the third site the length of the flowering season determined the reproductive output.     

Phenotypic plasticity and population differentiation in seeds and seedlings of the grass Anthoxanthum odoratum

Summary Seeds of Anthoxanthum odoratum were transplanted reciprocally between a xeric and a mesic field population that were genetically differentiated in adult traits. In one experiment seeds were reciprocally buried in bags in the soil, in a second experiment seeds were reciprocally sown in small plots. For most traits, site effects were much larger than seed-source effects. Germination, emergence, mortality of buried seed and recruitment were significantly higher at the xeric site than at the mesic site, irrespective of population of origin. Seed dormancy, was significantly higher for seed originating from the mesic than from the xeric population. Seedling recruits originating from the xeric population tended to be larger at both sites. Fecundity of seedling recruits depended on the environment; fecundities of plants growing in the xeric site had more than double the fecundity of plants growing in the mesic site. Phenotypic plasticity rather than population differences determined variation in performance in the seed and seedling stages.     

Ecological and evolutionary importance of neighbors in the grass Anthoxanthum odoratum

Summary A field experiment was performed to estimate the relative importance of neighbors and the rest of the environment for the growth, mortality and reproductive output of cloned individuals of the perennial bunchgrass Anthoxanthum odoratum. Single cloned Anthoxanthum tillers (targets) were reciprocally transplanted between a xeric and a mesic grassland site with one of four neighbor treatments: (1) no neighbors, (2) Anthoxanthum neighbors transplanted from the xeric site, (3) Anthoxanthum neighbors from the mesic site, and (4) Holcus lanatus neighbors. Targets without neighbors had a twofold higher two year reproductive output (RO) than those with neighbors, but there was no difference among neighbor treatments. No overall site effect on two year RO was found, because the site with the highest mortality among targets produced larger plants, with more inflorescences. Neighborhood competition was more intense at the xeric site than at the mesic site. The effects of environmental and neighborhood variation on Anthoxanthum were additive, rather than interactive. Population origin did not affect target performance significantly. Anthoxanthum neighbors of different origin did respond differentially to transplant site. There was a strong target genotype x site interaction, but no genotype x neighborhood interaction.     

Survival and recruitment favored by safe site-strategy – the case of the high alpine, non-clonal cushions of Eritrichium nanum (Boraginaceae)

In 14 permanent plots at four distant sites in the European Alps life tables, cohorts and recruitment of Eritrichium nanum were studied during a period of 5 resp. 6 years. The germination rates were tested in the Botanical Institute of the University of Basel and besides the life tables of the plots at natural habitats we followed the early stages of development in pot cultures at the University Botanical Garden in Basel. Most of the seedlings of E. nanum emerged in spring, in the pot cultures as well as under natural conditions, a behavior which confirms general findings. But the germination rates (0–30%) were markedly low. The pot cultures show that some seeds are after 4 or 5 years still viable. Premature flowering in the second summer is a rather common behavior of E. nanum seedlings, which happened in all the cohorts observed during five years, partly up to 20% of the total of living plants in the corresponding plots. It is evident that life expectancy of E. nanum plants increase rapidly with rising size, to such a degree that cushions of only 10–20 cm² surface have a survival chance of more than 70%, those of 20–30 cm² even of more than 95% during a period of 5 years. But it has to be mentioned that juvenile plants are not growing parallel to their lifetime, many of them persist during several years in the size categories I/II (0–5 cm²), while others suddenly arise from category I to III (–10 cm²) or even IV (–20 cm²), what is mostly caused by joining, a specific type of safe site strategy. Though not obligatory, juvenile plants growing close together, often add their sizes to form one larger cushion, which was found to be particularly advantageous for recruitment. This is obviously a primary strategy of E. nanum and favors, especially on bare ground, the survival of its descendants which was found higher than expected (average of 26% after 5 years’ growth at natural habitats). Focusing on the whole life cycle of E. nanum, the present studies show constraints in pollination, a small seed production and low germination rates to be well compensated by the high survivorship and successful recruitment of the seedlings.     

Variable demography in relation to germination time in the annual plant Tagetes micrantha Cav. (Asteraceae)

This study tests whether variability in a life history trait such as emergence time has ecological and evolutionary significance in seasonal environments due to their relationship with fitness components. The population dynamics of nine different cohorts of seedlings emergence were analyzed in a natural population of the annual herb Tagetes micrantha Cav. (Asteraceae). Temporal variation in seedling emergence was related to the pattern of precipitation, while spatial variation could be related to topographical heterogeneity. Plants that emerged at the beginning of the season (early germination) had lower probabilities of survivorship than those that emerged at the end of the season (late germination). In contrast, plants that emerged early had, on average, higher fecundity than those that emerged late. The net result of these contrasting patterns of survival and fecundity constitute a life history trade-off in T. micrantha.     

Ecological genetics of Bromus tectorum

Summary By incorporating demographic analyses of fitness components (e.g., survival and reproduction) within a reciprocal sowing design, we tested for 3 consecutive years whether local adaptation has occurred in the alien grass Bromus tectorum (cheatgrass) within 7 habitats along an environmental gradient from arid steppe to subalpine forest in the Intermontain Region of western North America. Patterns of emergence and survival were strongly influenced by the local environment. In terms of survival, expression of significant local adaptation in Tsuga heterophylla habitat varied among years. In contrast, relative differences in flowering time among seed sources were stable across sites and years. Populations from the arid steppe were the earliest to flower; flowering was latest in populations from the mesic Tsuga heterophylla habitat. In terms of net reproductive rate, evidence for local adaptation in B. tectorum was obtained in populations from habitats representing environmental extremes: an arid, saline site dominated by the shrub Sarcobatus vermiculatus and clearings within the cool, mesic Tsuga heterophylla forest habitat. Unlike the plants introduced from other sites, members of the resident population at the Sarcobatus site flowered and produced seeds before soil water became limiting. In contrast, net reproductive rates in other habitats were sometimes the lowest for populations in their home site. This lack of an advantage for local populations within more environmentally moderate sites suggests that limited dispersal may restrict the rate at which superior genotypes are introduced into a particular site.     

Sudden changes in environmental conditions do not increase invasion risk in grassland

After direct habitat transformation, biological invasions are considered to be the second most important threat to biodiversity. A better understanding of the factors affecting invasion success in new areas is crucial, and may provide insight into potential control actions. We hypothesized that invasion risk increases in habitats undergoing a sudden change in the disturbance regime or environmental conditions. For testing this assumption we initiated a seed sowing experiment while introducing two novel treatments, mowing twice and fertilizer application, in two grassland sites (one dryer and one mesic) in Romania. The seeds of two invasive species, Solidago canadensis and Rudbeckia laciniata, and two resident natives of similar seed sizes, life-forms and strategies were sowed in treated and control plots, and seed germination, seedling establishment and growth were followed during four months. Contrary to our expectations, there was no difference in the treatment effects on seed germination and seedling establishment between species, while there was on seedling vigour of the larger seeded species in the dryer grassland site, where the native had a higher performance especially in increased nutrient conditions. Indifferently from applied treatments, invasive species had greater cumulative germination in the mesic site, while natives were far more successful in seedling establishment in the drier site. At the same time, seed size was found to be a very important factor explaining germination and establishment success, with large seeded species outperforming small seeded species in any circumstances. Our results call the attention upon management interventions in mesic, productive grassland sites opening colonization windows for the recruitment of those invasive species of which ecological requirements correspond to local environmental conditions.     

Identification of conservation measures to protect the Japanese endangered plant species Aster kantoensis

To identify the factors responsible for degrading the habitat of the endangered plant species Aster kantoensis, as well as the vulnerable life stage where this occurs, we carried out sowing experiments. Two natural habitats were simulated, being situated along the floodplains of the Tama River in central Japan. Seeds collected from a natural habitat were sown in two apparently suitable locations (Tomoda and Ishida sites). Germination, survival, growth, and seed production were subsequently monitored from 1993 through to 1997. The Tomoda site was a gravel bar in floodplains formed by flooding in 1991, while the Ishida site (two plots) was one gravel bar where several plants were growing sparsely and another where a population had become extinct in 1992. Seed cohorts completed their life cycle within 3 years at the Ishida site and within 5 years at the Tomoda site. Monitored parameters at Ishida were substantially lower than those at Tomoda. In addition, estimates of population growth indicated an increase at Tomoda and a rapid decrease at Ishida. However, degradation of habitats seemed to occur at Tomoda over the monitored periods. In view of our results, we conclude that natural germination of about 0.13% is needed for increasing population size. The major factors for decreasing population size are believed to be the lack of safe sites for germination and seedling establishment in old habitats (>10years). Conservation measures are suggested based on these findings.     

On strict and facultative biennials

Summary Most so-called biennial plants may flower in their second growing season, but usually take longer in the field. Work reported here and in four recent papers on six different species has identified natural populations of strictly biennial plants, which flower only in their second year. The strict biennials and facultative biennials seem to form two distinct groups. All six strict biennial species were growing in infertile sites, and at least four of the six have annual variants at other sites, so they may be regarded as extended annuals growing in sites too infertile for the life cycle to be completed in one year. In contrast most facultative biennials are pauciennials (short-lived monocarpic perennials) of more fertile, disturbed habitats. Moreover, the rigid two-year life cycle of the strict biennials creates at each site two temporally separated sub-populations, which flower in odd and even years.     

Flowering requirements of Polymnia canadensis (Asteraceae) and their influence on its life history variation

The purpose of this study is to determine the flowering requirements of Polymnia canadensis and how they correspond to the occurrence of winter annuals, biennials, and short-lived monocarpic perennials in this species. Polymnia canadensis has a vernalization requirement for flowering, and even very small plants (i.e., those with one pair of leaves) can be vernalized. Vernalized plants can flower under both long- and short days. However, to flower plants must attain a minimum postvernalization size. Plants of this primarily monocarpic species that do not die after they flower once require another period of vernalization to flower a second time (i.e., to be dicarpic). Vernalized plants exposed to high temperatures can be devernalized; these must be re-vernalized in order to flower.     

Testing for local adaptation in Avena barbata: a classic example of ecotypic divergence

Forty years ago, Robert Allard and colleagues documented that the slender wild oat, Avena barbata , occurred in California as two multi-locus allozyme genotypes, associated with mesic and xeric habitats. This is arguably the first example of ecotypes identified by molecular techniques. Despite widespread citation, however, the inference of local adaptation of these ecotypes rested primarily on the allozyme pattern. This study tests for local adaptation of these ecotypes using reciprocal transplant and quantitative trait locus (QTL) mapping techniques. Both ecotypes and 188 recombinant inbred lines (RILs) derived from a cross between them were grown in common garden plots established at two sites representative of the environments in which the ecotypes were first described. Across four growing seasons at each site, three observations consistently emerged. First, despite significant genotype by environment interaction, the mesic ecotype consistently showed higher lifetime reproductive success across all years and sites. Second, the RILs showed no evidence of a trade-off in performance across sites or years, and fitness was positively correlated across environments. Third, at QTL affecting lifetime reproductive success, selection favoured the same allele in all environments. None of these observations are consistent with local adaptation but suggest that a single genotype is selectively favoured at both moist and dry sites. I propose an alternative hypothesis that A. barbata may be an example of contemporary evolution – whereby the favoured genotype is spreading and increasing in frequency – rather than local adaptation.     

Accumulation of buried seeds and establishment of ruderal therophytic communities in disturbed habitat,central Japan

The establishment and maintenance mechanisms of pioneer communities were investigated in ruderal habitats under two disturbance regimes, frequent and infrequent cutting sites. In the infrequent cutting sites, large perennials dominated through the year and inhibited the invasion of annuals, and the perennial community succeeded to forest stage if the cutting was stopped. In the frequent cutting sites, therophytic communities of winter and summer annuals alternated by season. Fresh seeds of both winter and summer annuals are dormant, but they have different germination times and thus can share the same sites in different seasons. Wind-dispersed biennials and large perennials have nondormant seeds and easily invade the sites; however, they are unable to mature to reproductive phase due to recurrent cuttings. The therophytic species, which can complete their life-cycle in a period between cuttings, accumulate seeds in the soils and are maintained by these buried seeds (seed bank annuals) during recurrent disturbances. The seed bank is compensation for the dispersal inefficiency of seed bank annuals. After abandonment of the frequent cutting sites, the buried seeds of seed bank annuals germinate and become the first-year pioneers. Thus, seed bank annuals are not invasive colonizers but are the remnants of the ruderal weed communities before abandonment.     

Invasive species and climate change: <Emphasis Type="Italic">Conyza canadensis</Emphasis> (L.) Cronquist as a tool for assessing the invasibility of natural plant communities along an aridity gradient

The predicted reduction in precipitation in the eastern Mediterranean due to climate change may expose the natural plant communities to invasive species. We assessed whether natural plant communities along an aridity gradient in Israel were resistant to invasion by considering differences in abiotic conditions and community characteristics in these regions. We considered Conyza canadensis as a model plant as it is a common invader in the region. We examined the mechanisms and functional traits of both the plant communities and C. canadensis that promote or discourage invasion. Study sites represented a rainfall gradient with four ecosystem types: mesic Mediterranean, Mediterranean, semiarid and arid. Our results showed that the mechanisms of community invasion resistance varied along the aridity gradient. At the arid and semiarid sites, water deficiency impaired the establishment of C. canadensis. At the mesic Mediterranean site, plant competition had a negative effect on C. canadensis performance, thus greatly reducing the likelihood of its establishment. We conclude that a decrease in regional precipitation due to climate change may not affect intrinsic resistance characteristics of natural plant communities to invasion in the area.     

Eco‐evolutionary responses of Bromus tectorum to climate change: implications for biological invasions

How plant populations, communities, and ecosystems respond to climate change is a critical focus in ecology today. The responses of introduced species may be especially rapid. Current models that incorporate temperature and precipitation suggest that future Bromus tectorum invasion risk is low for the Colorado Plateau. With a field warming experiment at two sites in southeastern Utah, we tested this prediction over 4 years, measuring B. tectorum phenology, biomass, and reproduction. In a complimentary greenhouse study, we assessed whether changes in field B. tectorum biomass and reproductive output influence offspring performance. We found that following a wet winter and early spring, the timing of spring growth initiation, flowering, and summer senescence all advanced in warmed plots at both field sites and the shift in phenology was progressively larger with greater warming. Earlier green‐up and development was associated with increases in B. tectorum biomass and reproductive output, likely due early spring growth, when soil moisture was not limiting, and a lengthened growing season. Seeds collected from plants grown in warmed plots had higher biomass and germination rates and lower mortality than seeds from ambient plots. However, in the following two dry years, we observed no differences in phenology between warmed and ambient plots. In addition, warming had a generally negative effect on B. tectorum biomass and reproduction in dry years and this negative effect was significant in the plots that received the highest warming treatment. In contrast to models that predict negative responses of B. tectorum to warmer climate on the Colorado Plateau, the effects of warming were more nuanced, relied on background climate, and differed between the two field sites. Our results highlight the importance of considering the interacting effects of temperature, precipitation, and site‐specific characteristics such as soil texture, on plant demography and have direct implications for B. tectorum invasion dynamics on the Colorado Plateau.     

Phenotypic plasticity and interpopulation differences in life history traits of<Emphasis Type="Italic"> Armadillidium vulgare</Emphasis> (Isopoda:Oniscidae)

The hypothesis that the balance of trade-offs between survivorship, growth and reproductive allocation in the terrestrial isopod Armadillidium vulgare will change when resource input is increased has been investigated experimentally. When the quality of food available was increased, by adding a mixture of litter from herbaceous dicotyledonous plants to a background low-quality food of dead grasses, survivorship was found to be the most phenotypically plastic trait, increasing by 168%. Growth rates increased by 99% but reproductive allocation by only 21%. In the field, members of a population from a site with more high-quality food grew more than twice as fast as those from a site where less high-quality food was available. The population from the site with higher food availability, contrary to predictions from the laboratory study, did not survive as well as that from the site with less available high-quality food. This may be because the site that is more favourable for growth has a more stressful physical environment due to much bigger temperature fluctuations, which are known to be an important cause of mortality in this species. When individuals from both populations were reared under controlled laboratory conditions, both the parental and F1 generations from the poor growth environment survived better than those from the good growth habitat. However, even when given an excess of high-quality food those from the poor growth environment continued to grow more slowly and had a lower reproductive allocation than those from the site with higher food availability. We conclude that microevolutionary changes may have occurred in the balance of resource allocation between survivorship, growth and reproductive allocation, to favour higher survivorship during the longer prereproductive period at the site where growth to the threshold size for reproduction takes longer.     

Variation in radial growth responses to drought among species, site, and canopy strata

 Radial growth responses to drought were examined in the tree-ring records of six species growing within two locations of differing land-use history and soil moisture characteristics, and in overstory and understory canopy positions in northern Virginia. Tree species experienced differential ring-width reductions during or immediately following four severe drought periods occurring from 1930 to 1965 and were influenced by climatic variables including annual and summer temperatures, annual precipitation, and annual Palmer Drought Severity Index. Relative growth comparisons averaged across species before and after drought years indicated that understory trees on dry-mesic sites grew 11% faster after drought compared to pre-drought rates while mesic site trees in both canopy positions grew approximately 4% slower. Superposed epoch analysis indicated that Liriodendron tulipifera growing on mesic sites experienced greater ring-width reductions associated with drought than co-occurring, more drought-tolerant Quercus alba and Q. velutina. On dry-mesic sites, L. tulipifera also experienced greatly reduced growth as a result of drought but exhibited significant growth increases following individual drought events. Quercus alba was the only species that exhibited a consistent, significant ring-width decrease associated with all droughts on dry-mesic sites. In contrast, Pinus virginiana was least impacted by drought on dry-mesic sites but was much more impacted by drought on mesic sites, indicating a drought×site interaction for this species. Overstory Carya glabra and Q. alba experienced larger growth decreases during drought on dry-mesic versus mesic sites. Understory tree growth reductions did not differ between site types but were often significantly larger than overstory responses of the same species on mesic sites. Following drought, most trees exhibited growth reductions lasting 2–3 years, although several species experienced reductions lasting up to 6 years. The results of this study suggest that tree rings represent an important long-term proxy for leaf-level ecophysiological measurements of growth responses to drought periods. Received: 31 July 1996 / Accepted: 16 April 1997     

Drought response and changing mean sensitivity of European beech close to the dry distribution limit

European beech (Fagus sylvatica L.) covers a large area mainly in the colline and montane ranges in Europe, and a drier and warmer climate, as expected for the coming decades, is likely to alter its distribution. So far, an altitudinal shift has been projected using a variety of modelling approaches. However, we lack knowledge about the climatic and edaphic factors that control the growth and competitive behaviour of beech at its dry distribution limit. We applied and further developed dendroecological methods to study the drought response and sensitivity pattern of beech at sites with different moisture regimes. We compared three pairs of sites from different geographical regions near the dry distribution limit of beech in Switzerland, consisting of a dry and mesic site each. Radial growth differed between mesic and dry sites, in that average ring-width at mesic sites was around double the width at dry sites. For the whole study period (1930–2006), the sites with the lowest available soil water capacity (AWC) were found to respond most sensitively to drought. However, in recent years, sites with higher AWC have shown increasing drought sensitivity, i.e. they have responded even more strongly to drought than the dry sites. This change in sensitivity corresponds to a seasonal shift in drought response at mesic sites, with a change in the months showing significant drought response in all three studied regions compared with the past. Even though dry sites generally displayed a larger number of negative pointer years than mesic sites, it appears that the frequency of pointer years has increased at mesic sites, i.e. they have become more sensitive particularly in the last quarter of the twentieth century. Yet, the frequency of pointer years at the dry sites has remained fairly constant. These results indicate that beech trees near their dry distribution limit are adapted to extreme conditions already, while changes in the growth patterns of beech under mesic conditions have to be expected.     

Shoot demography in new England populations of Maianthemum canadense desf.

Summary The demography of shoots of eight populations of a herbaceous perennial exhibiting clonal growth, is presented. The study was done along an elevational gradient, from a more open secondary mixed forest to a denser, more mature stand. Most shoots lived one to three years on the average, but shoots as old as twelve years were found. Large variation in formation and mortality of shoots was observed among plots and years. Yearly trends in the mortality rates of site replicates showed a higher correlation than rates of shoot formation. Although the density of shoots was highest in the drier sites, the turnover of shoots was highly variable and apparently uncorrelated with site location. Age structures revealed a tendency of longer-lived and higher reproductive activities among shoots from more mesic sites. It is hypothesized that environmental rather than density controls are primary causes of the population dynamics observed in this species.