EFFECTS OF LIGHT AND DENSITY ON RESOURCE ALLOCATION IN A FOREST HERB,ASTER ACUMINATUS (COMPOSITAE)

Recent studies show that forest herbs have low sexual reproductive efforts (SRE) and high vegetative reproductive efforts (VRE) and that a variety of factors may affect the levels of reproduction and the distribution patterns of forest species. Aster acuminatus is a micro-successional forest herb common in northern New England. Resource allocation patterns were determined for seven aster patches varying in density and light level. Plant density affects neither plant size nor any aspect of resource allocation, and VRE remains constant over all density and light levels. Light level is, however, significantly correlated with patch location, average plant size and average patch SRE. Only large plants produce flowers suggesting that light affects sexual reproduction indirectly through effects on plant size. The high rhizome and low seed survivorship of this species may partly explain the constant VRE and varying SRE levels, although the resource allocation patterns observed do not correspond well with the predictions of certain theoretical models.     

The developmental ecology of mycorrhizal associations in mayapple, Podophyllum peltatum, Berberidaceae

Associations between plants and arbuscular mycorrhizal (AM) fungi are widespread and well-studied. Yet little is known about the pattern of association between clonal plants and AM fungi. Here we report on the pattern of mycorrhizal association within the rhizome systems of mayapple, Podophyllum peltatum. Mayapple is a long-lived understory clonal herb that is classified as obligately mycorrhizal. We found that while all mayapple rhizome systems maintained mycorrhizal associations, the percent colonization of roots by AM fungi differed among ramets of different age. The highest concentrations of AM fungi were in the roots of intermediate-aged ramets, while roots beneath the youngest ramet were not colonized. This pattern of ramet age or position-dependent colonization was observed in two separate studies; each conducted in a different year and at a different site. The pattern of AM fungal colonization of mayapple rhizome systems suggests that the mycorrhizal relationship is facultative at the ramet level. This conclusion is reinforced by our observation that augmentation of soil phosphate lowers root colonization by AM fungi. We also found that soil phosphate concentrations were depleted by ca. 1% under the same ramet positions where roots bore the highest AM fungal loads. Three non-exclusive hypotheses are proposed regarding the mechanisms that might cause this developmentally dependent pattern of mycorrhizal association.     

THE RELATIONSHIPS BETWEEN SEASONAL VARIATION IN LIGHT INTENSITY,RAMET SIZE,AND SEXUAL REPRODUCTION IN NATURAL AND EXPERIMENTAL POPULATIONS OF ASTER ACUMINATUS (COMPOSITAE)

We conducted experiments to evaluate the effects of different patterns of seasonal light availability on ramet size and sexual reproduction in the understory herb, Aster acuminatus. In April 1982, 200 potted rhizomes were divided equally between a high light and a shaded site. Twice during the growing season subsets of pots were reciprocally transferred between the sites, resulting in eight different light treatments. There was significant variation among treatments in mean ramet weight (0.41–1.21 g), height (15.0–23.4 cm), number of flowers (0.4–7.9) and percentage of ramets flowering (14–95). Periods of high light increased ramet height, weight, rhizome production and flowering, but high light earlier in the season increased ramet height more than did high light later in the season, while high light later had pronounced effects on the other three traits. Measurements taken in nine natural patches experiencing different patterns of seasonal light availability are consistent with the experimental results. The results provide new insights into the general phenomenon of size-dependent reproduction in herbs and illustrate how the phenology of biomass allocation can affect growth and reproduction.     

Light-induced variation in the growth and dynamics of transplanted ramets of the understory herb,Aster acuminatus

Summary Ramets of the understory herb, Aster acuminatus, were transplanted from two source populations into eight understory garden sites that varied in light and soil moisture levels. Ramet growth, clonal growth, flowering and survivorship were monitored for three growing seasons. Large differences among gardens in ramet growth, clonal growth and flowering developed in the first growing season and increased in the next two years. This variation was positively correlated with garden light level but not at all with soil moisture. Mortality rates were low in all gardens and showed that genets from any particular source could survive over a broad range of environmental conditions. There was no conclusive evidence for any source population differences in the capacity to survive or grow in different environments. The rapid, light-induced responses of transplanted ramets resulted in garden populations very similar in appearance to natural populations experiencing similar light regimes. These results combined with those from other studies of A. acuminatus provide strong evidence for the importance of light in explaining population patterns and dynamics in this species.     

Reproductive traits in the tropical moss Octoblepharum albidum Hedw. differ between rainforest and coastal sites

Abstract

Octoblepharum albidum Hedw. is an autoicous moss commonly occurring in tropical savannas, dry forests, rainforests and coastal habitats. It frequently reproduces by spores and asexual structures (gemmae and protonemata or buds at leaf tips), making it a good model for understanding how reproductive traits change with respect to habitat type. Our aims were to characterize the different life-history traits in O. albidum relative to sexual and asexual cycles and to detect variations in reproductive performance among the different habitats, trade-offs among these traits, and relationships among reproductive traits and plant length. We studied colonies from two Atlantic rainforests and two coastal sites in north-eastern Brazil. Shoots in the coastal sites, compared to those of the forest sites, had higher numbers of sporophytes, male and female branches per shoot, male gametangia per sexual branch, and longer setae. Numbers of female gametangia per sexual branch did not differ between forest and coastal sites. A male-biased sex ratio of branches and gametangia occurred in all sites. Compared to gemmae, sporophytes and protonemata or buds were more likely to be found on longer shoots than on shorter ones, but this relationship was only applicable to forest sites. The abundant production of gemmae and protonemata or buds at leaf tips, and sporophytes (spores) in O. albidum are important components in explaining colonization success and maintenance in this tropical moss. Longer sporophytic setae in addition to a higher reproductive performance (especially for number of male gametangia and sporophytes per shoot) may favour spore dispersal and colonization in plants of coastal sites.     

Fragmentation in Montipora ramosa: the genet and ramet concept applied to a reef coral

The fates of Montipora ramosa fragments have been followed and related to physical factors on the reef. The degree of genetic homogeneity between colonies of M. ramosa was assessed by reciprocal grafting techniques, and the cross reef cline interpreted in terms of the degree of fragmentation. Two botanical terms, genet and ramet, referring to individuals derived from sexual (planula settlement) and asexual (fragmentation of colonies) processes respectively have been introduced to the coral literature.     

Translocation of Photoassimilates Between Sister Ramets in Two Rhizomatous Forest Herbs

Autoradiography and liquid scintillation techniques were usedto trace the pattern of photoassimilate translocation in twoperennial forest herbs, Aster acuminatus Michx. and Clintoniaborealis (Ait.) Raf. Vegetative shoots of C. borealis emergeearly each spring from the growing apices of a long-lived rhizomesystem. Vegetative shoots of A. acuminatus emerge in late springfrom rhizomes that decay within 2 years. In both species rametssurvive for only one growing season. Any connected plants aresisters. Mature leaves of these two species were exposed for1 h periods to ¹⁴CO₂ during spring and summer. Radioactivitycould subsequently be detected in exposed leaves (16–84per cent, depending on the season), adjacent above ground plantparts (0.7–23 per cent), roots and rhizomes (4.9–84per cent) and, when present, flowers (4–8.5 per cent).Old rhizomes of C. borealis are prominent storage sites forphotosynthate. In A. acuminatus, no significant translocationbetween sister ramets (i.e. above ground shoots connected bya common rhizome) was observed. In C. borealis, there was small,but consistent translocation between sister ramets (0.2–4per cent). Disturbance of unexposed sister ramets by defoliation,shading or herbivory increased the flow of photoassimilatesto disturbed parts in C. borealis, but not in A. acuminatus.Based on the absence of translocation flow, ramets of A. acuminatusmay be regarded as physiologically independent. Connected rametsof C. borealis show physiological integration. These resultsare correlated with ecological differences between the two species. Aster acuminatus, Clintonia borealis, translocation, ramet, vegetative reproduction, forest herb     

Forests are not immune to plant invasions: phenotypic plasticity and local adaptation allow <Emphasis Type="Italic">Prunella vulgaris</Emphasis> to colonize a temperate evergreen rainforest

In the South American temperate evergreen rainforest (Valdivian forest), invasive plants are mainly restricted to open sites, being rare in the shaded understory. This is consistent with the notion of closed-canopy forests as communities relatively resistant to plant invasions. However, alien plants able to develop shade tolerance could be a threat to this unique forest. Phenotypic plasticity and local adaptation are two mechanisms enhancing invasiveness. Phenotypic plasticity can promote local adaptation by facilitating the establishment and persistence of invasive species in novel environments. We investigated the role of these processes in the recent colonization of Valdivian forest understory by the perennial alien herb Prunella vulgaris from nearby populations in open sites. Using reciprocal transplants, we found local adaptation between populations. Field data showed that the shade environment selected for taller plants and greater specific leaf areas. We found population differentiation and within-population genetic variation in both mean values and reaction norms to light variation of several ecophysiological traits in common gardens from seeds collected in sun and shade populations. The colonization of the forest resulted in a reduction of plastic responses to light variation, which is consistent with the occurrence of genetic assimilation and suggests that P. vulgaris individuals adapted to the shade have reduced probabilities to return to open sites. All results taken together confirm the potential for rapid evolution of shade tolerance in P. vulgaris and suggest that this alien species may pose a threat to the native understory flora of Valdivian forest.     

Age structure in Liatris acidota (Compositae)

Summary Ten populations of the perennial herb, Liatris acidota Engelm. & Gray (Compositae), growing on the Texas Gulf Coast (USA) were analyzed for age structure, frequency of predation, and frequency of asexual reproduction. Nine of the ten populations exhibited non-stable age distributions due to a deficiency of young non-seedling plants. Age structure was not related to levels of predation nor to the amount of asexual reproduction. Environmental perturdation is suggested as the cause of the observed deviation from a stable age distribution.     

A comparison of plant communities on the basis of their clonal growth patterns

This study was performed to analyse how a vegetative propagation pattern of plants affects the coexistence of species and subsequent species richness of the community. We compared community average clonal growth in the herbal communities of forests, wooded meadows, and open meadows in Laelatu, Estonia. The parameters used for the calculation of the community averages and measured for each species were ramet life span, rhizome branching, and clonal mobility. We also examined the intrinsic (i.e. independent of the environment) relationship between community clonal growth and plant species density. We found strong correlations between the environmental factors (productivity, light availability, and mowing regime) and community averages of clonal growth parameters, while species density was (negatively) correlated only with community average of rhizome increment. The community average of ramet life span decreased with the increasing biomass of the herb layer. No evidence was found to support the hypothesis that species-rich communities may consist of species with more contrasting mobility compared with species-poor communities. Independent of the effect of the environmental factors, species density was positively correlated with ramet density. There was intrinsic positive relationship between species density and community average of ramet life span at open meadow sites and intrinsic negative relationship between species density and community average of rhizome increment at wooded meadow sites. We conclude that in forest communities the capability of clonal plants to forage for light is favoured, while in unmown meadows a competitively strong phalanx growth form is advantageous. We established that ramet turnover increases and vegetative mobility decreases with increasing species diversity, although these two relationships depend strongly on the type of the studied community.Co-ordinating editor: J. Tuomi     

Presidential Address—Some Reflections On The Nature Of Species

Background: Variation in the distribution and abundance of woody plants as consequence of disturbances such as fire may be explained by lineage age.

Aims: We tested whether lowland tropical tree lineages that colonise secondary forests are more late-diverging than clades from old-growth forests, and whether tree phylogenetic beta diversity from old-growth to secondary forests is higher in burned than non-burned secondary forests.

Methods: We sampled tree communities in old-growth forests and in secondary forests with distinct disturbance histories (burned and unburned). We calculated mean family age in each plot, and tested for differences among forest types using ANOVA. A phylogenetic fuzzy-weighting procedure was employed to generate a matrix describing the abundance of tree clades per plot, which was then analysed using a principal coordinate analysis.

Results: Most clades found in old-growth forests were underrepresented in secondary forests, which have been heavily colonised by a single species from a young lineage that is not found in old-growth forests. Phylogenetic beta diversity was higher between unburned secondary forests and old-growth forests than between burned secondary forests and old-growth forests.

Conclusions: The capacity of Neotropical trees to colonise secondary forests and persist after fire disturbance may be related to the age of distinct lineages.     

Anthropogenic disturbances and plant biodiversity in forests of Uttaranchal,central Himalaya

Eight forest types varying in disturbance frequencies were identified along an elevational gradient in Uttaranchal, central Himalaya. Low elevation forests were close to human habitation and had high disturbance frequency, while high elevation forests were situated far from the human habitation and had low disturbance. The dominant tree species at low elevation were Pinus roxburghii and Quercus leucotrichophora, while Q. floribunda and Q. semecarpifolia dominated the high elevation forests. Pyracantha crenulata was the shrub present in all the forests except in Q. semecarpifolia forest and Anaphalis contorta, a herb species, was present in all the forests. Disturbance decreased the dominance of single species and increased the plant biodiversity by mixing species of different successional status. Species richness and diversity for all the vegetation layers were higher in low elevation–high disturbance forests. Mean tree density decreased from high to moderate and increased in low disturbance. The shrub density decreased from high to low disturbance while the reverse occured for herbs. High proportion of early successional species in disturbed forests indicated that disturbance induces succession. The mean number of young individuals increasing from high to low disturbance indicates that disturbance adversely affects regeneration. But, however, the high number of young individuals of Coriaria nepalensis, a small non-leguminous nitrogen fixing tree, in disturbed forests shows that the forest is regenerating. This species could be helpful in the re-establishment of original vegetation through triggering the regeneration of these forests. High elevation–low disturbed forests separated from low elevation–high disturbed forests. Forest type and elevation may have more influence on tree richness while shrub and herb richness may be more sensitive to disturbance and forest types.     

贵州百里杜鹃风景区旅游干扰对杜鹃群落特征的影响

为揭示旅游干扰对于百里杜鹃风景区杜鹃群落特征影响,选取杜鹃纯林和杜鹃混交林两种杜鹃群落,分析干扰对群落结构及物种多样性的影响。采用典型抽样的方法进行样地设置和调查,通过旅游影响系数分析样地旅游干扰等级,并分析对比了不同旅游干扰程度和干扰方式对群落特征的影响。研究表明：1）百里杜鹃景区共有植被67科102属130种,其中杜鹃纯林共有植被97种,以马缨杜鹃、箭竹和露珠杜鹃为优势物种,杜鹃混交林共有植被115种,以马缨杜鹃、露珠杜鹃和映山红为优势种;2）两种杜鹃群落均在中等干扰水平时具有最高的Berger-Parker丰富度指数和Shannon多样性指数,中度干扰一定程度上促进了杜鹃群落物种多样性增加,不同的是两种群落的Shannon指数变化趋势不同,在杜鹃纯林中表现为中度干扰>轻度干扰>重度干扰,而在杜鹃混交林中中度干扰>重度干扰>轻度干扰,相较于杜鹃纯林,杜鹃混交林物种更为丰富,在旅游干扰下生物多样性更为稳定,抗干扰能力更强,而杜鹃纯林受旅游干扰波动显著;3）垃圾指数、踩踏指数与杜鹃混交林物种多样性指数、群落生长结构无明显相关性,而垃圾指数与杜鹃纯林Shannon多样性指数、物种数以及草本均高呈显著负相关,踩踏指数与杜鹃纯林灌木层高度、盖度呈现极显著负相关,与草本均高呈显著正相关;4）各杜鹃群落对旅游干扰有不同响应,为保持杜鹃群落稳定性,应在花季适当分流,严格控制杜鹃纯林的重度干扰。     

Sexual dimorphism in clonal growth forms and ramet distribution patterns in <Emphasis Type="Italic">Rumex acetosella</Emphasis> (Polygonaceae)

We investigated clonal traits in the dioecious herb Rumex acetosella to characterize sexual dimorphism in clonal forms and to correlate below-ground clonal patterns and above-ground ramet distributions. We recorded creeping root length, branching patterns, ramet and clump (caespitose ramets from the same position on the root) sprouting patterns, and biomass allocations in three females and males. We also estimated the patch size of flowering ramets within a quadrat. No sexual dimorphism was detected in the frequencies of branches and flowering ramets per root length. Male plants allocated proportionally more biomass to below-ground organs. Total root length did not differ between the sexes. Females sprouted more clumps with fewer flowering ramets per root length than males, which sprouted fewer clumps with more flowering ramets, which meant that clump sprouting patterns were phalanx-like in females and guerrilla-like in males. Flowering ramets were aggregately distributed in both females and males and patch sizes were similar between sexes, indicating that the spreader propagations were not found in the guerrilla-like males. We assumed that sexual dimorphism occurred in response to physiological integration for higher reproductive effort in females.     

Experimental analysis of mating patterns in a clonal plant reveals contrasting modes of self‐pollination

Hermaphrodite plants commonly practice self‐fertilization (selfing), but the mechanisms responsible vary depending on the mode of self‐pollination, pollinator behavior, and degree of clonality. Whether selfing occurs within (autogamy) or between flowers (geitonogamy) is of evolutionary significance because their fitness consequences differ. We used floral manipulations and genetic markers to determine the relative contribution of autogamy and within‐ versus between‐ramet geitonogamy to the selfing rate of the bumblebee‐pollinated, clonal herb Aconitum kusnezoffii. Data on flowering phenology and bumblebee foraging were also collected to determine opportunities for different modes of self‐pollination. Autogamy accounted for only 12% of the selfing rate with the remainder resulting from geitonogamy. Whole‐ramet emasculation of clones with multiple ramets reduced selfing by 78%, indicating that within‐ramet geitonogamy contributed significantly (68%) to total selfing. Selfing of single‐ramet plants was 44% less than multiple‐ramet plants, indicating that the contribution of between‐ramet geitonogamy was substantially less (20%) than within‐ramet geitonogamy, probably because of bumblebee foraging behavior. Our results demonstrate for the first time in a clonal plant that within‐ramet geitonogamy is substantially greater than between‐ramet geitonogamy and highlight the importance of considering the influence of clonal architecture and pollinator foraging on modes of self‐pollination.     

Maintaining diversity through intermediate disturbances: evidence from rodents colonizing rehabilitating coastal dunes

Rodents inhabit the coastal dune forests of KwaZulu‐Natal, South Africa. Here habitat rehabilitation following mining of dunes has resulted in coastal dune forest succession similar to that recorded in nonmined forests. We investigated the colonization of rehabilitating stands and evaluate the role of disturbance in maintaining rodent diversity. A trapping programme was established between July 1993 and February 1995 during which rodent colonization, local extinction and species richness were recorded for rehabilitating stands of different ages. Trends in these variables were closely associated with one of three possible outcomes for a disturbed patch over time, with no intervening disturbances following the initial disturbance. Colonization was initially high which led to an increase in species richness. Extinction was lower than colonization, but became higher when the habitat was 3 years old, which led to a decline in richness. We extrapolate this result assuming negligibly small disturbances after the initiation of rehabilitation and suggest that intermediate levels of disturbance maintain rodent species richness in coastal dune forests. Furthermore, our results illustrated species turnover, a prediction of the recorded outcome, with young stands dominated by Mastomys natalensis and older stands by Saccostomus campestris or Aethomys chrysophilus.     

Dances with fire: Tracking metapopulation dynamics of<Emphasis Type="Italic">polygonella basiramia</Emphasis> in Florida scrub (USA)

The regional persistence of species subject to local population colonization and extinction necessarily depends on how landscape features and disturbance affect metapopulation dynamics. Here, we characterize the metapopulation structure and short-term dynamics ofPolygonella basiramia. This rare, short-lived perennial herb is endemic to Florida scrublands and lacks a seed bank. Fires create the open sand gaps within a shrub matrix that support this species but also kill established plants. Thus, persistence depends on frequent colonization of unoccupied gaps. We are monitoring population dynamics within and among 1204 gaps distributed among 19 shrub patches. Considerable subpopulation turnover is evident at the gap level with rates of gap extinction exceeding rates of colonization in the first year. Whether declines in overall abundance continue is likely to depend on patterns of disturbance and regional stochasticity in this dynamic landscape.Polygonella is more likely to occupy larger and less isolated gaps, demonstrating that landscape features and disturbance strongly affect metapopulation dynamics. BecausePolygonella basiramia displays characteristics, occupancy patterns, and turnover dynamics consistent with metapopulation theory, it represents a model system for studying plant metapopulations.     

Are boreal ecosystems susceptible to alien plant invasion? Evidence from protected areas

Although biological invasion by alien species is a major contributor to loss of indigenous biological diversity, few studies have examined the susceptibility of the boreal biome to invasion. Based on studies of other ecosystems, we hypothesized that alien plants will be restricted to disturbed areas near human activity and will not be found in natural areas of boreal ecosystems in Gros Morne National Park (Canada), a protected area experiencing a wide range of disturbance regimes. The distribution of alien plants in the region was evaluated using surveys, and study sites were established in naturally and anthropogenically disturbed habitats that had been invaded. Within study sites, randomization tests evaluated the importance of disturbance to alien plant invasion by examining changes in environmental conditions and species abundance within various disturbance regimes, while the importance of site characteristics limiting the distribution of alien plants were examined using Canonical Correspondence Analysis. Consistent with studies in a variety of biomes, areas of high disturbance and human activity had the greatest abundance of resources and the highest percentage of alien species. However, contrary to our hypothesis, natural areas of boreal ecosystems were found susceptible to alien plant invasion. Vegetation types vulnerable to invasion include forests, riparian areas, fens, and alpine meadows. Natural disturbance occurring in these vegetation types caused increases in bare ground and/or light availability facilitating alien plant invasion. Although high soil pH was associated with alien plants in these areas, disturbance was not found to cause changes in soil pH, suggesting susceptibility to invasion is pre-determined by bedrock geology or other factors influencing soil pH. Moose (Alces alces), a non-native herbivore, acts as the primary conduit for alien plant invasion in GMNP by dispersing propagules and creating or prolonging disturbance by trampling and browsing vegetation. The recurrent nature of disturbance within the boreal biome and its interaction with site conditions and herbivores enables alien plants to persist away from areas of high human activity. Managers of natural lands should monitor such interactions to decrease the invasion potential of alien plants.     

Disturbances by human trampling alter the performance,sexual reproduction and genetic diversity in a clonal woodland herb

Disturbances play a major role in structuring the community composition in a variety of ecosystems. The potential impact of disturbances on sexual reproduction and genetic diversity of plants has so far received little attention. We examined the effects of disturbance by human trampling on the performance, sexual reproduction and clonal diversity in the woodland herb Anemone nemorosa in six sites disturbed by intensive recreational activities and in six undisturbed sites in suburban forests in Switzerland. Disturbance by human trampling decreased shoot height of A. nemorosa and altered the size-dependent flowering probability function. Furthermore, in disturbed sites an increased percentage of aborted seeds was found, resulting in a reduced sexual reproductive potential of A. nemorosa populations. Both clonal and genetic diversity of A. nemorosa were reduced in disturbed sites. Our study shows for the first time that disturbance by human trampling can decrease the genetic diversity in a clonal woodland plant. This suggests that genetic aspects of woodland plants should be considered in restoring degenerated forest sites.     

Environmental limitation contributes to the differential colonization capacity of two forest herbs

Question: The recovery of forest plant communities in post‐agricultural landscapes is largely determined by dispersal constraints, but can environmental legacies of former land use additionally limit the recolonization of recent forests by forest herbs? Location: Ancient forest and recent forest on former heavily fertilized agricultural land (Muizenbos, northern Belgium). Methods: Seeds and adults of two forest herbs with similar life‐history traits, but contrasting colonization capacity – the fast‐colonizing Geum urbanum and the slow‐colonizing Primula elatior– were introduced into both ancient and recent forest sites. Soil conditions and plant tissue nutrient concentrations were measured to characterize habitat quality. To determine whether the introduced species could successfully establish and persist, we monitored recruitment, longevity and adult performance during 8 years in permanently marked plots. Results: Phosphorus availability was ten times higher in recent forest soils and was also reflected in the plant tissue samples. Species longevity was clearly lower in recent forest sites indicating higher turnover. The fast‐colonizing G. urbanum counterbalanced this lower longevity by new establishment, while the slow‐colonizing P. elatior dropped below the number of originally introduced individuals. Additionally, G. urbanum performed better in recent forest sites in contrast to P. elatior. Conclusions: Even when dispersal constraints of the slow‐colonizing forest herb P. elatior are eliminated through introduction, environmental conditions in recent forest sites additionally restrict its recruitment, longevity and performance. These experimental results suggest that environmental constraints may strengthen the differences in colonization capacity among forest herbs if slow dispersers also tend to be less likely to establish.