Life-history monographs of Japanese plants 14: Cardamine leucantha (Tausch) O. E. Schulz (Brassicaceae)

The life-history characteristics of Cardamine leucantha (Tausch) O. E. Schulz (Brassicaceae) are described. The species is an herbaceous perennial that favors open but relatively moist habitats. It is distributed from Kyushu to Hokkaido in Japan but also occurs in Korea, Mongolia, China and the Russian Far East. In southwestern Japan, shoots start sprouting from mid- to late April, reaching approximately 30–70 cm in height, with 5–10 compound leaves. Ramets simultaneously produce one or more stoloniferous rhizomes that elongate until new ramets are formed at the tips. Cardamine leucantha has a pseudo-annual life cycle, in which mother ramets wither at the end of each season and only daughter ramets appear aboveground in the next year. As a result, ramet positions change annually. In a study population, the number of flowers averaged 23.9 ± 21.0 per ramet and fruit set was 44.2 ± 24.8% (10.4 ± 10.1 fruits per ramet). Ramets produced 3.8 ± 2.3 rhizomes that were 22.0 ± 15.6 cm long. The species sometimes forms large populations. A single genet develops into a group of disconnected ramets spreading via clonal growth. Reproductive characteristics (e.g., fruit set and numbers of flowers and rhizomes) vary among populations, resulting in interpopulation differences in genet structure. Because the reference genome became available recently, established molecular tools will be applied effectively for the investigations of C. leucantha as a model clonal plant.     

扎龙湿地芦苇分株生态可塑性及其对土壤因子的响应

扎龙湿地的芦苇既可形成大面积的单优群落,也可形成不同群落斑块。采用大样本抽样调查与统计分析方法,对湿地内水生生境、湿生生境、旱生生境和盐碱生境芦苇种群分株高度和生物量进行比较。结果表明,6—10月份,4个生境芦苇种群分株高度及生物量均以水生生境最高,盐碱生境最低,水生生境株高为盐碱生境的1.5—2.3倍,分株生物量为2.0—5.1倍,生境间的差异性以及差异序位均相对稳定。4个生境株高生境间变异系数(19.45%—31.56%)均高于生境内变异系数(8.07%—17.61%),分株高度在生境间的可塑性更大;分株生物量中水生生境、湿生生境和盐碱生境3个生境间的变异系数(33.43%—55.61%)均低于生境内变异系数(44.85%—79.82%),分株生物量在生境内的可塑性更大。不同生境条件下芦苇种群分株,在生长和生产上均存在较大的生态可塑性,表现出明显的环境效应,其中土壤含水量是该地区芦苇分株生态可塑性变异的主要驱动因子(R0.80),为正向驱动。     

The demography of a clonal herbaceous perennial plant, the longleaved sundew Drosera intermedia, in different heathland habitats

Eight populations of longleaved sundew Drosera intermedia Hayne ramets were studied in three hydrologically and phytosociologically differing heathland habitats over a 4 yr period a pool edge, an old path through the wet heath, a seepage area At the extreme sites, with high levels of water table fluctuations (pool edge and dry path), both survival and recruitment were low, resulting in small populations The pool edge populations went extinct At the less extreme sites (wet paths and seepage area), populations were stable, but ramet and genet turnover was higher at the path sites than in the seepage area Demographic variation between populations was explained by differences in adult survival and sexual recruitment (asexual recruitment did not differ clearly), which, in turn, were related to specific biotic and abiotic factors Likely adult mortality causes were summer drought (all habitats), winter frost (path habitat), cover with a crust of iron bacteria after heavy summer rainfall (seepage area) and shading by Sphagnum and algae (pool edge) Low seedhng recruitment was caused by a low germination succes (pool, seepage area) and seedling survival (seepage area, dry path)     

Ramet performance in two tussock plants — Do the tussock-level parameters matter?

Studies in plant demography are primarily done at the level of ramets and typically collect ramet-related parameters such as ramet size, type and history. This approach ignores possible effects of factors associated with higher levels, such as genet or tussock. This is particularly important in perennial resprouting herbs with persistent root that consists of many ramets as interaction between ramets, both by competition and by resource sharing, are likely to be intense in these plants. This study investigates effects of tussock-level parameters (age, size and ramet position within tussock and ramet density) on performance of individual ramets in two tussock-forming resprouting herbs (Tanacetum vulgare and Centaurea jacea).The results show that position of a ramet within tussock did not affect ramet growth, but had significant effects on flowering and survival in both species. The direction of the effect differed between the two species; marginal ramets were more successful in T. vulgare, while central ramets were more successful in C. jacea. In addition, tussock age had a significant effect on ramet flowering and survival in T. vulgare. Both these effects are likely to be due to the more competitive life form of T. vulgare, which is restricted to temporary habitats with intense competition. C. jacea is a species of mown or grazed grasslands with lower productivity, which are more stable and where competition is weaker. The effects of the number of neighboring ramets and of the tussock size were significant, but often locality-specific.The results indicate that although many important effects of tussock-related parameters exist, their direction and magnitude differ between species or even populations and are thus not easily predictable. Neglecting these effects, however, is likely to make the demographic models weaker.     

A predictive relationship between population and genetic sex ratios in clonal species

Sexual reproduction depends on mate availability that is reflected by local sex ratios. In species where both sexes can clonally expand, the population sex ratio describes the proportion of males, including clonally derived individuals (ramets) in addition to sexually produced individuals (genets). In contrast to population sex ratio that accounts for the overall abundance of the sexes, the genetic sex ratio reflects the relative abundance of genetically unique mates, which is critical in predicting effective population size but is difficult to estimate in the field. While an intuitive positive relationship between population (ramet) sex ratio and genetic (genet) sex ratio is expected, an explicit relationship is unknown. In this study, we determined a mathematical expression in the form of a hyperbola that encompasses a linear to a nonlinear positive relationship between ramet and genet sex ratios. As expected when both sexes clonally have equal number of ramets per genet both sex ratios are identical, and thus ramet sex ratio becomes a linear function of genet sex ratio. Conversely, if sex differences in ramet number occur, this mathematical relationship becomes nonlinear and a discrepancy between the sex ratios amplifies from extreme sex ratios values towards intermediate values. We evaluated our predictions with empirical data that simultaneously quantified ramet and genet sex ratios in populations of several species. We found that the data support the predicted positive nonlinear relationship, indicating sex differences in ramet number across populations. However, some data may also fit the null model, which suggests that sex differences in ramet number were not extensive, or the number of populations was too small to capture the curvature of the nonlinear relationship. Data with lack of fit suggest the presence of factors capable of weakening the positive relationship between the sex ratios. Advantages of this model include predicting genet sex ratio using population sex ratios given known sex differences in ramet number, and detecting sex differences in ramet number among populations.     

扎龙湿地不同生境芦苇分株的形态特征及生物量分配的异速关系

不同环境条件下的植物个体可以表现出形态特征的分异和物质分配的权衡与调整。采用大样本抽样调查与统计分析方法,比较研究扎龙湿地不同生境芦苇(Phragmites Australis)生殖株和营养株的形态特征以及生物量分配的异速关系。结果表明：在9月末,盐碱生境、旱生生境、湿生生境和水生生境芦苇分株的生长表现出较大的生态可塑性,株高和株重均以盐碱生境最小,水生生境最大,最大值与最小值的比值分别为1.3—3.3和1.8—5.1,分株生长在种群间的变异度高于种群内,与营养株相比,生殖株的变异度较低;分株的支持分配与生产分配的比值为1.8—4.2,生产分配以盐碱生境最高,以水生生境最低,而支持分配和生殖分配表现与生产分配相反的序位;生殖株的花序长和花序重与株高间呈直线函数形式增长,株高和株重低于种群平均值的20%和35%的分株不进行有性生殖;叶重、叶鞘和茎重以及分株重与株高间呈幂函数形式的异速生长关系。植物通过改变个体的形态特征以及调整构件间生物量分配适应不同环境,而受遗传因素控制的构件间生长关系却相对稳定。     

Habitat and sex specific differences in the dioecious weed Acetosella vulgaris (Polygonaceae)

Abstract Patterns of resource allocation, numbers of reproductive structures and sex ratios of flowering populations of the dioecious weed Acetosella vulgaris (Fourr.) were examined in the Kosciuszko alpine region of Australia. Specifically, the sex‐specific response of ramets was compared between a disturbed alpine habitat, in which weeds such as A. vulgaris are common (disturbed roadside/path‐edge), and a native alpine habitat in which weeds appear to have a limited capacity to germinate, grow and reproduce (undisturbed tall alpine herbfield). The disturbed habitat was generally more favourable for growth of A. vulgaris. Ramets of both sexes had greater total and vegetative biomass than ramets in native habitats. Although reproductive biomass was also greater in disturbed habitats, females had less reproductive biomass than males, which was not as predicted. Reproductive effort was not affected by habitat or gender. The disturbed habitat also favoured increased numbers of inflorescences per ramet and flowers per ramet, as expected. Whereas the gender of the ramet also influenced numbers of reproductive structures, again, this was not as predicted. Females had more flowers per ramet and more flowers per inflorescence than males. This may be because of factors associated with wind pollination. Females were taller in native habitats but there was no difference between the sexes in disturbed habitats. Sex ratios varied from all male populations to nearly all female populations among the 25 sites sampled irrespective of habitat. Factors such as time since last disturbance may have contributed to variation in the sex ratios of alpine populations of A. vulgaris in Australia.     

扎龙湿地保护区异质生境芦苇种群分株构件的数量特征

采用单位面积取样,计数和测量的调查与统计方法,对扎龙湿地保护区4个生境单优群落芦苇种群分株构件数量特征进行比较分析。结果表明,4个生境芦苇种群从5月10日左右返青后进入营养生长期,分株高度和分株密度均持续增加到生殖生长初期的8月份,其中6-8月份差异均达到显著水平(P0.05),8-10月份差异均未达到显著水平(P0.05),芦苇进入生殖生长期后,分株便停止高度生长,地下芽的输出也不再形成分株补充现实种群;分株生物量和种群生物量均持续增加到生殖生长旺盛期的9月份,至休眠期的10月份均有所降低,各月份间的差异均达到显著水平,芦苇种群在生长季末期,具有将生产的物质分配转移到地下储藏与营养繁殖器官的形成与生长上的特性。芦苇种群分株数量特征与返青后实际生长时间之间均较好地符合对数函数关系,R~2在0.818-0.994之间,拟合方程均达到了P0.05的显著水平。4个生境芦苇种群分株密度以湿生生境最大,依次为水生生境、旱生生境,盐碱生境最小,分株高度、分株生物量和种群生物量均以水生生境最大,依次为湿生生境、旱生生境,盐碱生境最小。因此,4个生境芦苇种群分株构件数量特征均表现出基本一致的生育期节律性,同时,芦苇种群的个体生长和种群动态存在明显的环境效应,土壤含水量、pH是影响该地区芦苇分株数量特征的主要因子。     

Effects of soil heterogeneity and clonal integration on Alternanthera philoxeroides populations with a radial ramet aggregation

Some clonal plants can spread their ramet populations radially, and soil heterogeneity and clonal integration may greatly affect the establishment of these types of populations. We constructed Alternanthera philoxeroides populations with a radial ramet aggregation, allowing old ramets of clonal fragments to concentrate in central pots and younger ramets to root in peripheral pots. The peripheral pots were supplemented either with three different levels (high, medium and low) of soil nutrients to simulate a heterogeneous soil environment, or only one medium level of soil nutrients to simulate a homogeneous environment. Stolon connections between the central older ramets and the peripheral younger ramets were left intact or severed to test the effect of clonal integration. The maintenance of stolon connection could induce the division of labor between different‐aged ramets, by increasing the root investment in central ramets and the above‐ground growth in peripheral ramets. The maintenance of stolon connection could improve the growth of the central and peripheral ramets, clonal fragments and even the whole population. However, the positive consequence in peripheral ramets and whole fragments was only detected in the high‐nutrient patch of heterogeneous treatment. In sum, in the population with the radial ramet aggregation, clonal integration can play a key role in the rapid recruitment of young ramets of A. philoxeroides fragments, as well as the expansion of the whole population. The magnitude of clonal integration also became more obvious in the peripheral young ramets and whole fragments that experienced high‐nutrient patches.     

RAMET DYNAMICS FOR THE CLONAL SEAWEED PTEROCLADIELLA CAPILLACEA (RHODOPHYTA): A COMPARISON WITH CHONDRUS CRISPUS AND WITH MAZZAELLA CORNUCOPIAE (GIGARTINALES)

Little is known about the dynamics and the ecological interactions among ramets (fronds) from populations of clonal red seaweeds. Small ramets are very difficult to tag, so their growth cannot be monitored directly. The temporal variation of the relationship between stand biomass and ramet density offers information on ramet performance. We calculated this relationship for an intertidal population of Pterocladiella capillacea (Gmelin) Santelices et Hommersand (Gelidiales) from Baja California, Mexico. Biomass and density were positively correlated on an annual basis, indicating that biomass accumulated without involving self-thinning among ramets. This contrasts with nonclonal seaweeds, for which self-thinning among individuals occurs during growth, but agrees with other clonal red seaweeds, such as Chondrus crispus Stackhouse and Mazzaella cornucopiae (Postels et Ruprecht) Hommersand (both Gigartinales). The growth pattern for these members of the Gelidiales and of the Gigartinales holds despite differences in holdfast morphology and ramet branching degree and despite differences in the capacity of coalescence during early stages, known only for the Gigartinales. The positive slope for the dynamic biomass–density relationship, on a bilogarithmic scale, was statistically steeper for M. cornucopiae than for P. capillacea and for C. crispus. This suggests that the addition of new ramets during the growth season may be relatively more beneficial for biomass accumulation rates for M. cornucopiae. This would be expected for high-intertidal species subjected to strong abiotic stress, for which ramet crowding constitutes a key protection. Pterocladiella capillacea occurs at the mid-intertidal zone and C. crispus at the subtidal zone, so ramets would be relatively less important in that respect.     

Ramet demography and ecological attributes of the perennial river corridor plant Cnidium dubium (Schkuhr) Thell. (Apiaceae)

Cnidium dubium is a highly endangered, perennial river corridor plant in Central Europe. Here we summarise a 4-year field study of its ramet demography, growing in two different flood meadows in the Lower Havel River Valley in northeastern Germany, emphasising the influence of site-specific abiotic factors, mowing and the summer flood of 2002.We recorded 1658 ramets at densities between 2 and 180 ramets m⁻², which varied over area and time, indicating a high small-scale turnover of this species within grasslands. The half-life of the different ramet cohorts was estimated to range from 0.1 to 1.3 years, but single ramets of more than 4 years old were also found. Ramets are monocarpic: they propagate only once in their life. The highest probability of a vegetative ramet becoming a flowering stalk was 11.6%. Ramet density, cumulative number of newly produced ramets per year, leaf length, leaf age, reproductive growth and recovery from the summer flood of 2002 varied, depending on site. C. dubium had a higher performance at sites in intermediate zones of the overall flooding gradient compared with higher zones. Flooding stress is the controlling factor for the lower limit of its performance, while there was no evidence for an upper limit controlled by summer drought. Mowing slightly increased the cumulative number of leaves per ramet, but it decreased the size of new leaves and also the summer above-ground biomass per ramet. As a consequence, the most marked effect was that fewer ramets entered the generative life stage, but the plants were able to keep their overall reproductive performance on a level independent of mowing. Knowledge of basic growth patterns of C. dubium in natural populations could help to explain the species rareness, and may also suggest appropriate strategies to manage its survival.     

异质水分环境下野牛草相连分株间光合同化物的生理整合及其调控

异质环境下,克隆植物通过生理整合机制使资源在分株间实现共享,提高了其对异质性环境的适应能力,具有重要的生态进化意义,研究生理整合机制及其调控机理可为进一步发掘克隆植物应用潜力提供理论依据。以野牛草3个相连分株为材料,对其中一个分株用30%聚乙二醇6000(PEG-6000)模拟水分胁迫,通过Hoagland营养液培养试验,研究了异质水分环境下光合同化物在野牛草相连分株间的生理整合及分株叶片与根系内源激素ABA与IAA含量的变化规律。结果表明,14C-光合同化物在克隆片断内存在双向运输,但以向顶运输为主,异质水分环境下,受胁迫分株光合同化物的输出率明显降低,而与其相邻分株合成的光合同化物向受胁迫分株方向运输率明显增加;异质水分环境下,各分株ABA含量均明显增加,但以受胁迫的分株叶片及根系ABA的含量增加幅度最大,各分株IAA含量较对照均显著下降(P0.05),且以受胁迫分株IAA含量下降幅度最大;各分株叶片与根系ABA/IAA均显著提高(P0.05),相邻分株ABA/IAA增加幅度低于受胁迫分株。异质水分环境影响野牛草克隆分株间光合同化物的生理整合,且ABA与IAA在分株间光合同化物运输与分配过程中具有重要的调节作用。     

Assessing the importance of disturbance, site conditions, and the biotic barrier for dandelion invasion in an Alpine habitat

Several factors have been identified as relevant in determining the abundance of non-native invasive species. Nevertheless, the relative importance of these factors will vary depending on the invaded habitat and the characteristics of the invasive species. Due to their harsh environmental conditions and remoteness, high-alpine habitats are often considered to be at low risk of plant invasion. However, an increasing number of reports have shown the presence and spread of non-native plant species in alpine habitats; thus, it is important to study which factors control the invasion process in these harsh habitats. In this study, we assessed the role of disturbance, soil characteristics, biotic resistance and seed rain in the establishment and abundance of the non-native invasive species Taraxacum officinale (dandelion) in the Andes of central Chile. By focusing on human-disturbed patches, naturally disturbed patches, and undisturbed patches, we did not find that disturbance per se, or its origin, affected the establishment and abundance of T. officinale. The abundance of this non-native invasive species was not negatively related to the diversity of native species at local scales, indicating no biotic resistance to invasion; instead, some positive relationships were found. Our results indicate that propagule pressure (assessed by the seed rain) and the abiotic soil characteristics are the main factors related to the abundance of this non-native invasive species. Hence, in contrast to what has been found for more benign habitats, disturbance and biotic resistance have little influence on the invasibility of T. officinale in this high-alpine habitat.     

Clonal Diversity in Differently-Aged Populations of the Pseudo-Annual Clonal Plant Circaea lutetiana L.

Abstract: In many clonal plant species seedling recruitment is restricted to short colonization episodes early in the development of the population, and clonal diversity (i.e., genet diversity) in the population is expected to decrease with increasing population age. In established populations of the pseudo-annual Circaea lutetiana seedling recruitment has previously not been observed. Therefore, we expected established populations to have low clonal diversities. We analysed number and frequency of genets and spatial distribution of genets in six differently-aged C. lutetiana populations with the use of four informative RAPD primers. We found relatively low clonal diversities in young populations but very high clonal diversities in established populations. Therefore, the hypothesis was rejected that seedling recruitment does not occur in established populations. Moreover, we did not find large genet size asymmetries in established populations. Genet size differences can be caused by stochastic processes or by fitness related traits, such as differences in vegetative reproduction. Because vegetative propagation of ramets is dependent on ramet size, and the number of ramets and the size of each ramet determine genet size, we expected that large genets produced, on average, large ramets. However, this was not the case, suggesting that stochastic processes caused genet size differences. Genet size may also be bounded if spatial distribution of genets is affected by micro-habitat differences. For this we expected to find a clumped spatial distribution of ramets of the same genet. However, ramets of large genets were always found intermingled with ramets belonging to other genets.     

Effects of biotic and abiotic factors on the distribution of trout and salmon along a longitudinal stream gradient

Synopsis We examined the influence of biotic and abiotic factors on the distribution, abundance, and condition of salmonid fishes along a stream gradient. We observed a longitudinal change in fish distribution with native cutthroat trout, Oncorhynchus clarki utah, and introduced brown trout, Salmo trutta, demonstrating a distinct pattern of allopatry. Cutthroat trout dominated high elevation reaches, while reaches at lower elevations were dominated by brown trout. A transition zone between these populations was associated with lower total trout abundance, consistent changes in temperature and discharge, and differences in dietary preference. Variation in cutthroat trout abundance was best explained by a model including the abundance of brown trout and diel temperature, whereas variation in brown trout abundance was best explained by a model including the abundance of cutthroat trout and discharge. These results suggest the potential for condition-mediated competition between the two species. The results from our study can aid biologists in prioritizing conservation activities and in developing robust management strategies for cutthroat trout.     

Patterns of Solidago altissima ramet growth and mortality: the role of below-ground ramet connections

Summary For the rhizomatous perennial, Solidago altissima, I identified clonal fragments in the field, mapped ramet spatial locations, and documented patterns of ramet recruitment, growth, and mortality. Parent ramet size influenced the size and number of daughter ramets produced, and small ramets had lower survivorship and fecundity than large ramets. Similarly, small rhizomes tended to develop into small ramets, and ramets that survived to produce daughter ramets had longer parent-daughter rhizome connections than ramets that did not survive. In addition, most ramets that died during the growing season were connected to (genetically identical) ramets that persisted. There were large size inequalities among rhizomes, ramets, and clonal fragments. Inequalities in the size of ramets increased during the early part of the growing season, then decreased at the end of the season; similar patterns were observed for the growth of clonal fragments. In both instances, the decrease in size inequality could be attributed to the mortality of small individuals (ramets or clonal fragments). I found little evidence that ramet size hierarchies were structured by intraspecific competition. For example, path analyses and randomization tests indicated that size variation among S. altissima ramets was influenced little by the size of their near neighbors (but was influenced by parent size and rhizome size). In addition, within-season variation for the relative size and growth rate of individual ramets led to poor correlations between early and final ramet size; this result suggests that there was no stable hierarchy of dominant and suppressed ramets. I discuss implications of my results for contrasting interpretations of clonal plant population dynamics.     

Biogeography of parasitism in freshwater fish: spatial patterns in hot spots of infection

Contrary to species occurrence, little is known about the determinants of spatial patterns of intraspecific variation in abundance, particularly for parasitic organisms. In this study, we provide a multi‐faceted overview of spatial patterns in parasite abundance and examine several potential underlying processes. We first tested for a latitudinal gradient in local abundance of the regionally most common parasite species and whether these species achieve higher abundances at the same localities (shared hot spots of infection). Secondly, we tested whether intraspecific similarity in local abundance between sites follows a spatial distance decay pattern or is better explained by variation in extrinsic biotic and abiotic factors between localities related to local parasite transmission success. We examined the infection landscape of a model fish host system (common and upland bullies, genus Gobiomorphus: Eleotridae) across its entire distributional range. We applied general linear models to test the effect of latitude on each species local abundance independently, including the abundance of each co‐infecting species as another predictor. We computed multiple regressions on distance matrices among localities based on abundance of each of the four most common trematode species, as well as for geographic distance, biotic and abiotic distinctness of the localities. Our results showed that the most widely distributed parasites of bullies also achieve the highest mean local abundances, following the abundance – occupancy relationship. Variation in local abundance of any focal parasite species was independent of latitude, the abundance of co‐occurring species and spatial distance or disparity in biotic attributes between localities. For only one parasite species, similarity of abundance between sites covaried with the extent of abiotic differences between sites. The lack of association between hot spots of infection for co‐occurring species reinforces the geographic mosaic scenario in which hosts and parasites coevolve by suggesting non‐deterministic, species‐specific variation in parasite abundance across space.     

松嫩平原野古草种群构件结构动态

野古草是根茎型无性系禾草,在松嫩平原草甸经常形成单优种群落。采用单位面积挖掘取样、分株按营养繁殖世代划分龄级、根茎按实际生活年限划分龄级的方法,对松嫩平原单优群落和混生群落的野古草种群构件结构进行了调查与分析。结果表明,在生长季初期两群落野古草种群均以春性分株和根茎芽占优势,且分株及芽构件结构相对稳定,芽库的输出率单优群落为80.4%,混生群落为62.5%;整个生长季分株由2—3个龄级组成,1a分株数量是2a的2.9—10.2倍,其生物量各月份所占比例平均为93%,随着龄级的增加依次明显减少,呈增长型年龄结构;根茎由3—4个龄级组成,根茎累积长度及生物量均以2a占绝对优势,为稳定型年龄结构;分株生产力1a明显高于2a,对种群贡献最大;根茎贮藏力除个别月份以3a、4a最高外,两群落大部分以2a最高,在生长季后期,1a根茎物质积累的速率最快。     

Spatial distribution pattern of a clonal species: effects of differential production of clonal and sexual offspring

The spatial distribution patterns of genets and ramets within populations are expected to change as a function of the frequency with which clonal species recruit different types of offspring (sexual and clonal). We used an integrated approach to study the spatial arrangement of clonal plants by combining molecular and ecological data using Opuntia microdasys as a study system. The species is able to produce two types of clonal (plantlets and cladodes) and one type of sexual (seeds) offspring. Additionally it is found in three habitats that cause differences in the ability of each type of offspring to establish. In 2007, all individuals in the three habitats (162 in BH = bajada, 264 in IDH = hill-piedmont, and 136 at HPH = interdunes) were tagged and mapped. Amplified inter-simple sequence repeats (ISSR’s) were used to determine the multilocus genotype and relatedness of each individual ramet using 120 polymorphic bands (104 in BH, 128 in HPH and 180 in IDH). The spatial distribution pattern of genets and ramets was analyzed with the Hopkins test and spatial autocorrelation analysis. For all habitats we found that O. microdasys displayed a spatial distribution characterized by clumps of aggregated ramets, but habitats differed in the number of genets present. As for other clonal species a strong positive spatial autocorrelation exists within 20 m, although all analyses suggest that adjacent ramets are genetically less related to each other or belong to different genets, that is, ramets of different genets are intermingled. The spatial arrangement of genets and ramets in O. microdasys between habitats closely matches the frequency of establishment of each type of offspring (e.g. the more clonal areas are clumped groups of related individuals). These results confirm that in two habitats (BH and IDH) clonal recruitment had been more common than in the other habitat (HPH).     

Seed origin determines the range expansion of the clonal grass Elymus athericus

The recent invasion of clonal grasses to novel habitats poses a threat to biodiversity in various habitats. Elymus athericus, a clonal grass of north-western European salt marshes, is currently increasing in abundance and invading new habitats. In this study, we analyzed controlling factors for seedling establishment of E. athericus in frequently flooded low marsh habitats. Here, biotic and abiotic conditions are very different from the conditions of the parental sites with established populations higher up in the marsh. Hence, we hypothesized that seedling establishment at the expanding low marsh edge would depend on the parental origin (either through maternal effects or heritable local adaptation). We further hypothesized that seedling origin interacts with biotic factors such as herbivory and competition as well as with abiotic factors like inundation frequency. We tested the dependence of seedling survival, growth and vegetative reproduction on these factors in a factorial transplant experiment on Schiermonnikoog. Survival was high, with 77% of the planted seedling surviving until the end of the experiment. Biotic factors had a much stronger effect on seedling growth and mortality than parental origin and were independent of inundation. However, parental origin strongly interacted with herbivory and competition, with seedlings performing better under the conditions that resembled their parental site.We conclude that seedlings of E. athericus, a species that was previously thought to occur only in mid- to high marsh elevation, can establish at a frequently inundated low-marsh sites. Long term survival and further invasion will primarily depend on biotic factors in interaction with seed origin. Our results suggest that next to herbivory, limitation of seeds adapted to colonizing conditions is likely to slow down range expansion.