Intraspecific adoption and foster feeding of fledglings in the North Island robin

We sampled soils and vegetation within and outside two sheep and rabbit exclosures, fenced in 1979, on steep sunny and shady slopes at 770 m altitude on seasonally-dry pastoral steeplands. The vegetation of sunny aspects was characterised by higher floristic diversity, annual species, and low plant cover. Here the exotic grass Anthoxanthum odoratum dominated on grazed treatments, and the exotic forb Hieracium pilosella on ungrazed. Shady aspects supported fewer, and almost entirely perennial, species. Here Hieracium pilosella dominated grazed treatments, but co-dominated with the exotic forb H. praealtum and the native grass Festuca novae-zelandiae on ungrazed treatments. There was 43% more biomass in exclosures (P < 0.01). Most of the biomass difference (4285 kg/ha) was from greater root mass (2400 kg/ha). 1385 kg/ha of the difference was from herbage and the remainder (500 kg/ha) from litter. Exclosures had 50 to 100% more Ca, Mg, K and P in the biomass (P < 0.05), but the effect on soils was limited to significantly higher concentrations of total N (P < 0.05) and exchangeable Mg (P < 0.01) in 0-7.5 cm soils. We conclude that stopping grazing for 16 years on seasonally-dry steeplands results in greater plant cover, approximately double the biomass of standing vegetation, greater biomass in roots, and more biomass nutrients relative to grazed areas. However, it does not favour native species and has little effect on soil nutrients or soil carbon. Stopping grazing alone therefore cannot be regarded as a comprehensive short- or medium-term vegetation or soil rehabilitation option.     

Effects of intraspecific competition on size variation and reproductive allocation in a clonal plant

Clonal plants grow in diameter rather than height, and therefore competition among genets is likely to be symmetric and to result in smaller variation in size of genets than in non-clonal plants. Moreover, clonal plants can reproduce both sexually and vegetatively. We studied the effects of density on the size of rosettes and of clones, variation in the size of rosettes and of clones, and allocation to sexual and vegetative reproduction in the clonal herb Ranunculus reptans . We grew plants from an artificial population of R. reptans in 32 trays at two densities. After four months, differences in density were still apparent, although clones in the low-density treatment had on average 155% more rosettes and 227% more rooted rosettes than clones in the high-density treatment. The coefficient of variation of these measures of clone size was 15% and 83% higher, respectively, in the low-density treatment. This indicates that intraspecific competition among clones of R. reptans is symmetric and increases the effective population size. Rooted rosettes were larger and varied more in size in the low-density treatment. The relative allocation of the populations to sexual and to vegetative reproduction was 19% and 13% higher, respectively, in the high-density treatment. Moreover, seeds produced in the high-density treatment had a 24% higher mass and a 7% higher germination percentage. This suggests that with increasing density, allocation to sexual reproduction increases more than allocation to vegetative reproduction in R. reptans , which corresponds to the response of some other species with a spreading growth form but not of species with a compact growth form. We conclude that intraspecific competition is an important factor in the life-history evolution of R. reptans because intraspecific competition affects its clonal life-history traits and may affect evolutionary processes such as genetic drift and selection through its effect on the effective population size.     

Habitat characteristics,stage structure and reproduction of colline and montane populations of the threatened species <Emphasis Type="Italic">Arnica montana</Emphasis>

Arnica montana, a characteristic plant of nutrient-poor grasslands, has strongly declined during the last decades, particularly at lower altitudes. To gain insights into the underlying reasons for this decline, we recorded differences in size, stage structure and reproductive traits between 21 colline populations of A. montana in the Ardennes-Eifel region (280–600 m a.s.l.) and 10 montane populations in the Vosges Mountains (1,200 m a.s.l.). Community composition of the Violion caninae vegetation at colline sites and Nardion strictae vegetation at montane sites indicated that temperature and moisture were the main abiotic factors differentiating between low and high altitudinal relevées. The proportion of flowering rosettes decreased with altitude, indicating a shift from sexual reproduction to clonal growth. In contrast to expectation, Ellenberg values for nutrient availability as well as the proportion of young rosettes and population size did not differ between colline and montane populations. However, population size decreased with nutrient availability, indicating eutrophication as the reason for population decline. In small populations, plant density, the proportion of flowering rosettes, the number of flowerheads per rosette and the number of seeds per flowerhead were lower, indicating less suitable conditions. Thus, preventing further eutrophication in both colline and montane populations will be crucial for the conservation of the species. Reduced reproduction may not be important for the population dynamics of this clonal plant in the short term, but could affect genetic diversity and survival in the long term.     

Interaction between Some Pasture Species and 2

Possible control options are investigated for the introduced Hieracium weeds, particular problems in South Island high country, New Zealand. In a pot experiment regression of input to output ratios of above ground biomass over successive harvests, from binary mixtures was used to determine the competitive interaction between 13 pasture species and two Hieracium species, H. pilosella and H. praealtum, in a low fertility soil. Treatments also included a factorial of presence or absence of compartments separating root and shoots of species. Species differed in their mean growth rate, relative to Hieracium species. The ranking of mean growth rate relative to H. pilosella was Trifolium repens (best), Bromus inermis, Sanguisorba minor, Festuca novaezelandiae, F. rubra, Arrhenatherum elatius, Anthoxanthum odoratum, Lotus corniculatus, Hypochoeris radicata, Trifolium repens, T. hybridum, T. medium and Astragalus cicer. Shoot partitions decreased Hieracium's interaction with white clover while root partitions increased interaction with B. inermis. However, the rates were not related to the proportion of Hieracium in the mixtures, indicating a general lack of specific competitive effects against Hieracium.     

Factors predisposing short-tussock grasslands to

The effects of environment and management on the composition of short-tussock grasslands and the abundance of the invasive weed Hieracium pilosella were investigated in two small catchments. Species composition and site factors were recorded on a total of 182 plots and the management history of each catchment was reviewed. H. pilosella was present on >80% of all plots, but was at an early stage of invasion in one catchment (<5% cover) and dominant in the other (25% cover). Classification and ordination revealed strong between-catchment differences in community composition that reflected differences in environment (soil fertility and rainfall), disturbance history (animal populations and burning), and the stage of invasion by H. pilosella. In both catchments H. pilosella tended to be least abundant on the wettest, driest, and most fertile soils. However, such relationships were weak. Generalised additive models and regression showed that in the earlier stage of invasion individual site factors explained less than 20% of the variation in H. pilosella cover. Topographic position and slope (both indicative of soil moisture) were the most significant combined predictors, but together explained only 32% of the variation. In the later stage of invasion individual factors explained up to 33% of the variation. Topsoil sulphur, slope, and topsoil calcium were the most significant combined predictors, but together explained only 53% of the variation. Between-catchment comparisons highlighted the inter-related roles of environment, disturbance history, geographic location, availability of H. pilosella propagules, and stage of invasion in more fully explaining the abundance of H. pilosella. Of five models that have been proposed for Hieracium invasion, the: "grassland decline" model best incorporated the inter-related factors that influence spatial and temporal variation in H. pilosella abundance in the study area. This model concentrates on identifying predisposing and trigger factors that increase the likelihood of invasion and accounts for multiple causes and interactions by specifying five key factors that influence the ability of a plant species to invade existing vegetation: environment, disturbance, vegetation structure and composition, life history attributes of the invader, and the availability of invading propagules. The model potentially provides a comprehensive framework for evaluating the causes of Hieracium invasion, targeting research effort, and developing sustainable management strategies.     

Optimum reproduction and dispersal strategies of a clonal plant in a metapopulation: a simulation study with Hieracium pilosella

Clonal spread is favoured in many plants at the expense of seed production in order to expand rapidly into open habitats or to occupy space by forming dense patches. However, for the dynamics of a population in a patchy landscape seed dispersal remains important even for clonal plants. We used a spatially explicit individual-based metapopulation model to examine the consequences of two trade-offs in Hieracium pilosella L: first, between vegetative and sexual reproduction, and second, between short and far-distance dispersal of seeds. Our main question was, what are the environmental conditions that cause a mixed strategy of vegetative and sexual reproduction to be optimal. The model was parameterised with field data on local population dynamics of H. pilosella. Patch dynamics were given firstly by disturbance events that opened patches in a matrix of a clonal grass that were colonisable for H. pilosella, and secondly by the gradual disappearance of H. pilosella patches due to the expanding grass. Simulations revealed opposing selection pressures on traits determined by the two trade-offs. Vegetative reproduction is favoured by local dynamics, i.e. the need for maintenance and expansion of established populations, whereas seed production is favoured by the necessity to colonise empty habitats. Similar pressures act on the proportion of seeds dispersed over short and far distances. Optimum reproductive and dispersal strategies depended on habitat quality (determined by seedling establishment probability), the fraction of dispersed seeds, and the fraction of seeds lost on unsuitable ground. Under habitat conditions supporting moderate to low seedling establishment, between 20% and 40% of reproductive effort in H. pilosella should be devoted to sexual reproduction with at least 10% of the seeds dispersed over distances suitable to attain empty patches. We conclude that in a spatially heterogeneous landscape sexual seed production in a clonal plant is advantageous even at the expense of local vegetative growth.     

Neighbor effects on germination, survival, and growth in two arctic tundra plant communities

In relatively harsh environments such as arctic tundra, abiotic factors have traditionally been considered the primary determinants of community structure, overwhelming any effects of biotic interactions such as competition. Two common low arctic tundra types that differ in soil properties, moist acidic and moist non-acidic tussock tundra (MAT and MNT, respectively), occur in close proximity in northern Alaska. Several plant species occur in both communities with different relative abundance, while others are restricted to one. This study experimentally examined how neighboring vegetation affects germination, survival, and growth of species in these two communities that differ in soil pH, cation availability, and other characteristics. Germination of sown seeds was greater than background levels suggesting seed limitation may restrict recruitment of these clonal, perennial species. Germination of sown seeds was greater at both sites when both mosses and vascular plants had been removed compared to plots with intact vegetation. However, neighbors had almost no effect on survival and growth of adult transplants. Patterns of germination, survival and growth of several species differed depending on the community of origin and the community of destination of the seeds or transplanted adults. For example, transplants of the sedge Eriophorum vaginatum grew better if they were from MAT, and this species germinated better when sown at MNT. Although of relatively short duration (three growing seasons), this study suggests that biotic interactions may affect local species composition by restricting germination and establishment in these two communities, but have less of an effect on adult plants. Not surprisingly, site-specific abiotic conditions also exhibit control over species occurrence and relative abundance. Without disturbance to clear bare ground for recruitment of new individuals, these populations for the most part must rely on clonal growth to persist.     

Life history trade-offs in Amphibromus scabrivalvis (Poaceae): allocation to clonal growth,storage, and cleistogamous reproduction

Life history trade-offs among clonal growth, storage, and sexual reproduction were investigated in the perennial grass Amphibromus scabrivalvis in relation to soil nutrients. This species exhibits clonal growth by producing rhizomes and stores reserves in the form of basal corms; seeds are matured in cleistogamous spikelets on panicles enclosed within the leaf sheaths along each culm. Ten seed-derived genotypes (clones) were separated into 72 ramets and planted in the greenhouse. Control ramets received only water while the remainder received fertilizer applied every 2 wk. Twenty-four ramets were harvested per clone at 11, 20, and 26 wk. The dry mass of corms, rhizomes, roots, shoots, and seeds were recorded. Biomass partitioning to rhizomes provided a measure of carbon allocation to clonal growth, partitioning to corms provided a measure of allocation to storage, and partitioning to seeds provided a measure of allocation to sexual reproduction. Allocation to most organs was significantly influenced by clone identity; fertilizer significantly increased allocation to corms and seeds at 20 wk, but never affected rhizome allocation at any age. Corm allocation increased from 2% at 11 wk to 27% at 26 wk; rhizome allocation decreased from 10% at 11 wk to 3% at 26 wk. Significant negative relationships were detected for rhizome vs. seed and corm vs. rhizome allocation in fertilized clones at 20 wk. This suggests an age-dependent physiological life history trade-off between clonality and sexual reproduction and between clonality and storage. In contrast, a significant positive relationship was consistently noted for corm vs. seed allocation in fertilized and unfertilized clones at 20 and 26 wk. The absence of a trade-off between storage and sexual reproduction may indicate that these two processes are not necessarily mutually exclusive components of life history.     

Genetic Allee effects on performance, plasticity and developmental stability in a clonal plant

Negative effects of small population size on fitness, so-called Allee effects, may threaten population persistence even in intact habitat remnants. We studied genotypes of 14 isolated populations of the clonal plant Ranunculus reptans, for which molecular genetic (RAPD-) variability is higher for large than for small populations. In a competition-free greenhouse environment vegetative offspring of genotypes from large populations produced more rosettes and flowers, indicating higher fitness. Within-genotype coefficients of variation in performance traits, indicating developmental instability, were lower for genotypes from populations with higher RAPD-variability. In competition with a taller grass, we found relative reduction in leaf length less pronounced for plants from large populations, suggesting higher adaptive plasticity. Our experimental study of a plant with predominantly vegetative reproduction suggests, that negative genetic effects of recent habitat fragmentation, which so far rather were expected in plants with frequent sexual reproduction, are more severe and more common than previously acknowledged.     

Soil-Ph Declines and Organic-Carbon Increases under Hawkweed (

Changes in soil chemistry in relation to hawkweed (Hieracium pilosella) presence were determined at a site receiving less than 600 mm rainfall where hawkweed was colonising Pallic Soils (yellow-grey earths). pH was significantly lower (by 0.5 units) and organic carbon values were significantly higher (0.7% absolute, 40% relative) within hawkweed patches than in adjacent soil, but there was no significant difference in total nitrogen. These differences are attributed to the change from sparse vegetation dominated by ephemeral annual plant species to continuous vegetation dominated by a perennial.     

不同更新方式对林地植被生长及土壤性状的影响

本文研究天然次生林采伐迹地的三种更新方式对林地植被及土壤性状的影响。结果表明,炼山人工造林对林地植被和土壤有一定的破坏作用,是影响林木生长、导致物种丧失、引起林地水土流失和地力衰退的重要原因。不同林分物种丰富度为:天然更新 >人促更新 >炼山人工造林;林下植被生物量为:人促更新 >天然更新 >炼山人工造林。不炼山的天然更新和人促更新,有利于物种多样性保护、植被生长、林地土壤理化性状改善和土壤肥力提高,并具有更高的经济效益。     

Influence of pastoral management on plant biodiversity in a depleted short tussock grassland, Mackenzie Basin

This study investigated the effects of different management inputs (fertiliser and seed) and grazing patterns on plant biodiversity in a short tussock grassland with a strong Hieracium pilosella component. Cover abundance of vascular and non-vascular plants and environmental variables were measured in 32 10×10-m plots located in five blocks with different management treatments. Ordination of the floristic data separated the block with the highest management inputs from other blocks. Several adventive species were significantly more abundant in this block, while several native species were either absent or uncommon, but were significantly more abundant in other study blocks. H. pilosella was significantly more abundant in blocks with lower management inputs. Diversity was significantly higher in the block with the highest management inputs. The native tussock Poa colensoi had significantly greater cover abundance while Festuca novae-zelandiae tussocks were significantly taller in this block. Our results suggest that high management inputs reduce the abundance of H. pilosella and diversity of native species, but increase the abundance of other adventive species and the cover and vigour of native tussocks. Our results highlight an interesting management conundrum for short tussock grasslands. No-input management is likely to result in a decline in native biodiversity, as well as production values, as H. pilosella mats deplete soil nutrients and restrict regeneration of native species. However, input of fertiliser and adventive seeds to enhance production values, although resulting in an increase in the vigour and abundance of some native species (mainly tussocks) and a reduction in H. pilosella abundance, will also result in a decline in overall native species richness.     

Bottlenecks and spatiotemporal variation in the sexual reproduction pathway of perennial meadow plants

Sexual reproduction is important for the growth of populations and the maintenance of genetic diversity. Several steps are involved in the sexual reproduction pathway of plants: the production of flowers, the production of seeds and the establishment of seedlings from seeds. In this paper we quantify the relative importance and spatiotemporal variability of these different steps for four grassland perennials: Centaurea jacea, Cirsium dissectum, Hypochaeris radicata and Succisa pratensis. We compared undisturbed meadows with meadows where the top soil layer had been removed as a restoration measure. Data on the number of flower heads per flowering rosette, the numbers of flowers and seeds per flower head, and the seedling establishment probabilities per seed were collected by field observations and experiments in several sites and years. Combination of these data shows that H. radicata and S. pratensis have higher recruitment rates (1.9 and 3.3 seedlings per year per flowering rosette, respectively) than the more clonal C. dissectum and C. jacea (0.027 and 0.23, respectively). Seedling establishment is the major bottleneck for successful sexual reproduction in all species. Large losses also occurred due to failing seed set in C. dissectum. Comparison of the coefficients of variation per step in space and time revealed that spatiotemporal variability was largest in seedling establishment, followed closely by flower head production and seed set.     

松嫩平原不同生境条件下羊草无性系的生长规律

通过对栽培条件下实验无性系的跟踪调查和对羊草草甸水淹后恢复演替过程中天然无性系种群的随机取样调查,分析了不同生境条件下羊草无性系的生长规律.结果表明,在松嫩平原具有充分生长空间而没有种间竞争的风沙土上,两个实验羊草无性系的移植时间相差1个月,翌年生长季末期无性系的大小相差近1倍,整个生长季营养繁殖数量的增长规律均为指数函数.在羊草草甸水淹后恢复演替6年中,在羊草+杂类草群落、羊草+寸草苔群落,羊草无性系种群营养繁殖数量的增长规律均为幂函数.在具有种间竞争的碱化草甸土上,羊草无性系在水淹后恢复演替过程中仍可以通过指数形式迅速地增加无性系的子代数量,同时也迅速地扩展着无性系的生态位空间.     

Causes and consequences of extreme variation in reproductive strategy and vegetative growth among invasive populations of a clonal aquatic plant, <Emphasis Type="Italic">Butomus umbellatus</Emphasis> L. (Butomaceae)

The mode of reproduction may influence the spread of invasive species by affecting evolutionary potential and dispersal ability. We sampled 51 introduced North American populations of the clonal aquatic plant Butomus umbellatus L. (flowering rush) and found extreme variation in sexual fertility caused by polyploidy. Populations consisted of diploids that produced thousands of viable seeds or of sexually sterile triploids. Although a trade-off between sexual and clonal reproduction predicts that the sexual sterility of triploids would be compensated for by greater clonal reproduction, a greenhouse experiment involving eight diploid and 10 triploid populations showed that diploid plants not only invest substantially in sexual structures but also make hundreds of tiny clonal bulbils on both rhizomes and inflorescences. In contrast, triploids do not make bulbils and have very limited scope for clonal multiplication and dispersal. Diploid populations were more frequent than triploid populations, especially in the Great Lakes region. This is probably because of the difference between cytotypes in clonal rather than sexual reproduction, as genetic analyses indicate a general lack of sexual recruitment in North America. Although triploids were less common, they have a wider geographical distribution. This could be due to a greater ecological tolerance resulting from polyploidy. However, genetic evidence suggests that triploids have become widespread via their use in and escape from horticulture. North America is being colonized by two distinct forms of B. umbellatus that differ strongly in reproductive strategy as well as the vectors and pathways of invasion.     

Differential responses of germination and seedling establishment in populations of Tragopogon pratensis (Asteraceae)

The establishment phase is an important bottleneck in the life cycle of plants. It consists of two steps that are rarely separated, i.e., the germination of seeds and the establishment of seedlings. Here we report the results of two experiments in which we independently investigated germination and seedling establishment in the greenhouse, under different grass vegetation treatments representing different regeneration niches. Seeds of Tragopogon pratensis from six populations and two habitat types were studied, three from roadside verges and three from hayfields. Germination percentages and germination speed were higher for seeds from roadside verges than for seeds from hayfields, but were little affected by treatment. In contrast, seedling growth was much lower in the tall grass vegetation, than in the short grass and especially the bare soil treatment. Seedling sizes were generally similar for different populations and habitat types. Our results thus show that the two early steps in the establishment phase of plants may respond very differently to the micro-environment, and may have a different selection history. Insight into the ecology and evolution of life histories may require that germination and establishment are considered separately.     

A contribution to characterisation of genetic variation in some natural Polish populations of Elymus repens (L.) Gould and Elymus hispidus (Opiz) Melderis (Poaceae) as revealed by RAPD markers*

To determine the relative importance of clonal growth and sexual reproduction, the Randomly Amplified Polymorphic DNA (RAPD) method was used to study genetic diversity and clonal structure of six populations of Elymus repens and four populations of Elymus hispidus from Poland. These outbreeding species are virtually self‐sterile and form widely spreading and long‐lived rhizomes. Using 12 primers, a total of 150 unambiguous RAPD fragments were amplified and scored. Results of AMOVA showed no significant genetic distinction between morphologically distinguished varieties of E. repens and E. hispidus. E. repens had slightly higher intra‐specific genetic polymorphism than E. hispidus; the percentage of polymorphic bands per population ranged from 38 to 49 and from 19 to 38 respectively. Clonal diversity measured using the Simpson diversity index (D) indicated different contributions of clonal reproduction in particular populations of E. repens (D: 0.20–0.72). Populations of E. hispidus were dominated by one or a few clones, which were generally restricted to a single population (D: 0.00–0.22). RAPD revealed that most genetic diversity resided within populations of the two studied species, suggesting that, despite their clonal character, propagation by seeds contributes considerably to reproduction of E. repens and E. hispidus.     

Restoration and creation of freshwater wetlands using seed banks

The minimum information about a seed bank needed for a wetland restoration or creation project is a species list. There are two basic techniques for determining the composition of seed banks: (1) mechanical separation of seeds from a volume of soil and (2) germination of seeds from a volume of soil under appropriate environmental conditions. The latter method always gives biased results. It is best to collect as many random samples as possible when sampling a wetland seed bank. These can be combined as needed for processing. Field studies in India have demonstrated that vestigial seed banks can be used to re-establish a former vegetation type in a monsoonal wet-land that had become overgrown by a species of grass. In less than a year, 9 of 1 I species in the vestigial seed bank were found growing in areas cleared of the grass. Vestigial seed banks of drained prairie wetlands in the northcentral United States contained a few wetland species after 70 years, although species diversity and seed density declined significantly after 20 to 30 years of drainage and cultivation. In Florida, U.S.A., wetlands have been established in strip-mined areas using donor soils from existing wetlands. Newly established wetlands quickly developed a dense cover of vegetation, although this vegetation often lacked many desirable wetland species. Experimental studies of soil moisture conditions using a seed bank from the Delta Marsh, Canada, demonstrated that soil moisture affected both the total number of seeds, and the relative proportion of seeds of each species that germinated from a seed bank. The density of seedlings of emergent wetland species in the treatments was directly proportional to soil moisture, while that of terrestrial annuals was inversely proportional. Emergent species made up nearly 90% of the seedlings in the wettest treatment and 0% in the driest.From a paper presented at the Third International Wetlands Conference, 19–23 September, 1988, University of Rennes, France.     

Long-term control of species abundances in a dry grassland: a spatially explicit model

Abstract. The long-term spatio-temporal dynamics of a sparse dry-grassland community (Thymo-Festucetum) is investigated by a spatially explicit individual-based simulation model and by analytical model equations. The community (investigated over 15 yr in permanent plots) is characterized by a permanently low cover (30–50 %), mainly of the perennial tuft grass Festuca cinerea. Seedling establishment and the fate of juveniles are strongly dependent on weather conditions. The simulation programme focuses on the mechanism of clonal growth of grasses and the reproduction of tufts by fragmentation. Questions answered by the modelling approach were (1) which life-history features of the species are responsible for their persistence and for the low vegetation cover of the community and (2) what are the main mechanisms of species interactions. Different sets of simulation runs, together with the evaluation of the analytical models, show: (1) long-term persistence of the main species is possible only by a combination of sexual and clonal reproduction; the low cover is due to low germination rate, low mortality and limited growth of tufts; (2) the intra and interspecific control of the community is performed mainly via a reduction by already established individuals; (3) persistence of uncommon species relies on a diaspore buffer in, or around, the community (‘spatial mass effect’).