Ming Dong

Invasive alien plants in China: role of clonality and geographical origin

Biological invasions have become a significant threat to the global environment. Unfortunately, to date there is no consensus on invasion mechanisms and predictive models. Controversies range from whether we can reliably predict which species may become invasive to which species characteristics (e.g., life history, taxonomic groups, or geographic origin) contribute to the invasion processes. We examined 126 invasive alien plant species in China to understand the role of clonality and geographical origin in their invasion success. These species were categorized into three groups (I, II, III) based on their invasiveness in terms of current spatial occupation and the degree of damage to invaded habitats. Clonal plants consisted of almost half (44%) of the 126 invasive species studied, and consisted of 66% of 32 the most invasive alien plant species (Group I). There was a significant positive relationship between clonality and species invasiveness. A 68% of the 126 species studied originated in the continent of America (North and/or South America). These preliminary findings support that America is the primary geographical origin of invasive alien plant species in China and that clonality of the invasive plant species contributed significantly to the their invasiveness. The results suggest an urgent need at the global scale to investigate the mechanisms whereby plant clonal growth influences plant invasions, and the need for a focus at regional scale to examine factors affecting the exchange of invasive plant species between America and China.

Plasticity in morphology and biomass allocation in Cynodon dactylon, a grass species forming stolons and rhizomes

Stolons and rhizomes may be important for the placement of leaves and roots in new microhabitats (foraging), as well as for the storage of resources. In a species which forms both types of structures, such as Cynodon dactylon, both such functions could be combined in a single structure, or rhizomes and stolons could exert different functions. We examine these functions in C. dactylon, by investigating plasticity in morphology and biomass allocation in response to variation in light and nutrient availability. Both stolon and rhizome branching intensities were reduced in response to lower light and lower nutrient levels. Stolons and their internodes elongated greatly under lower light levels, but slightly shorter under lower nutrient levels (...)

Clonal plasticity in response to rhizome severing and heterogeneous resource supply in the rhizomatous grass Psammochloa villosa in an Inner Mongolian dune, China

In a field experiment, Psammochloa villosa plants were subjected to rhizome severing. Severing rhizomes reduced growth in the young, detached rhizome segments compared to the controls in terms of all measured clonal growth-related characters, i.e. number of rhizomes and shoots, total rhizome length and total number of rhizome nodes. In a container experiment, the control ramets received uniform water and nutrient supply but in heterogeneous treatments high and low levels of water and nutrient supply, respectively were established. The number of ramets, total rhizome length, dry weight per ramet and biomass allocation to the rhizome had higher values at high water and nutrient supply, while spacer length (length of rhizome between shoots) and rhizome internode length were not affected. The local response of ramets given low water supply was enhanced due to connection to a well watered parent ramet in terms of number of ramets, total rhizome length and dry weight per ramet. A remote effect was not observed in the other treatments or in the other measured characters.

Morphological responses to local light conditions in clonal herbs from contrasting habitats, and their modification due to physiological integration

Morphological responses to light and effects of physiological integration on local morphological responses are examined for Hydrocotyle vulgaris and Lamiastrum galeobdolon, stoloniferous herbs from open fenlands and forest understoreys, respectively. An assessment was made of whether these clonal herbs of similar morphology but from contrasting habitats show different foraging behaviour for light. In a garden experiment, the plants wer subjected to four levels of light availability, and to a split treatment in which the primary stolons grew along the border of patches of the two intermediate light levels. In this treatment the plant parts on opposite sides of the primary stolons were in contrasting light environments. Petiole extension was more responsive to light conditions in Hydrocotyle than in Lamiastrum, while the opposite was true for leaf area. Both species showed similar responses in stolon internode length and specific leaf area (SLA). Integration did not significantly modify local responses in stolon internode length in either species. Local responses in petiole length, leaf area and SLA of Hydrocotyle ramets were not significantly affected by physiological integration, except for the SLA of ramets in high light which was evened out by integration. In contrast, in Lamiastrum, local responses in petiole length, leaf area and SLA of many ramets in the shaded and/or light patch were significantly evened out by integration. As a result, interconnected ramets in patches of different light supply developed very different morphologies in Hydrocotyle, but not in Lamiastrum. The results indicate that the species differed in ramet morphological responses to light intensity as well as in effects of integration on local morphological responses, and suggest that species from different habitats show different foraging behaviour for light.

ON PLASTIC AND NON-PLASTIC VARIATION IN CLONAL PLANT MORPHOLOGY AND ITS ECOLOGICAL SIGNIFICANCE

Morphological plasticity in clonal plants has received wide attention because localized plastic changes in spacer length, branching intensity and branching angle may enable clonal plants to place ramets selectively in the more favourable microhabitats within a heterogeneous environment. These responses have been interpreted in terms of foraging behaviour.Studies of morphological plasticity in clonal plants are usually carried out with one or two genotypes of a species, or with material of unknown genetic origin. Based on the concept of phenotypic plasticity, it is argued that such studies do not reveal whether plasticity in a population can be modified by natural selection. In addition, responses are often evaluated at two environmental conditions only, which may underestimate plasticity. Hence, our information on the ecological and evolutionary significance of morphological plasticity in clonal plants is still very incomplete.Two examples are given to show that stolon internode and rhizome lengths may vary considerably within an individual plant. Only a minor part of this variation may be plastic, i.e. the variation is hardly changed by the environmental conditions to which the plants are subjected. Hence, non-plastic variation in clonal morphology may exceed the degree of morphological plasticity. The non-plastic variation seems to originate from species-specific patterns of stolon and rhizome development.Marked non-plastic variation may obscure the effects of morphological plasticity on the placement pattern of ramets in the field, suggesting that plasticity in clonal morphology may not be very effective in terms of foraging for favourable patches. Possible reasons for the low levels of plasticity of clonal spacers are discussed.

MORPHOLOGY AND GROWTH OF STOLONS AND RHIZOMES IN THREE CLONAL GRASSES, AS AFFECTED BY DIFFERENT LIGHT SUPPLY

In this paper, the hypothesis is tested that, in clonal grasses producing stolons and/or rhizomes, stolons always show a higher morphological plasticity than rhizomes in response to variation in light availability.Agrostis stolonifera (a stoloniferous grass),Holcus mollis (a rhizomatous grass) andCynodon dactylon (a grass forming both stolons and rhizomes), were grown in pots and subjected to three levels of light intensities. Both stolons and rhizomes branched more intensively under higher light levels. Irrespective of species, stolons consisted of longer internodes under lower light levels, while rhizome morphology did not respond significantly. Biomass partitioning to rhizomes was lower under lower light intensities while partitioning to stolons was not affected. Rhizomes usually had more dormant buds than did stolons. Our results suggest that stolons serve primarily as foraging organs for light, whereas the main function of rhizomes is storage of meristem and carbohydrates, irrespective of whether the grass species involved produces both rhizomes and stolons or only one type of spacer.

MORPHOLOGICAL RESPONSES TO NUTRIENT AVAILABILITY IN FOUR CLONAL HERBS

This paper examines morphological plasticity of clonal plants of contrasting habitats and of contrasting architectures in response to nutrient supply. The hypotheses were tested that plants from rich habitats possess greater plasticity in response to variation in resource supply than species from poor habitats, and that rhizomatous species are less plastic in their response than stoloniferous species. Two sympodial rhizomatous herbs (Carex flacca, C. hirta) and two monopodial stoloniferous herbs (Trifolium fragiferum, T. repens) were subjected to four levels of nutrient supply in a garden experiment. One of the two species of each genus (C. hirta, T. repens) is from fertile and the other from infertile habitats. We measured 1) whole plant characters: total plant dry weight, number of modules (product of a single apical meristem) and number of ramets; 2) ramet characters: ramet leaf area and ramet height; and 3) spacer characters: branches per module, length per module and length per module internode. All measured characters in the Trifolium species significantly responded to treatment: the values for all measured characters increased with higher levels of fertilization. The differences in plant characters between fertilization levels were larger in Trifolium repens than in T. fragiferum in terms of whole plant characters, ramet characters and stolon internode length. The two Carex species did not differ in their responses to treatment in terms of most characters measured. In ramet characters and in some whole plant characters the species from fertile habitats were more plastic than those from infertile habitats. In spacer characters this pattern was not found. Foraging could not be demonstrated unequivocally. Morphological plasticity in the stoloniferous (Trifolium) species was much larger than in the rhizomatous (Carex) species. This seems in accordance with a foremost storage function of rhizomes, as against a foremost explorative function of stolons.

Importance of clonal plants and plant species diversity in the Northeast China Transect

In plant communities, the internal (genet-level) control mechanisms on a spatio-temporal scale of clonal plants impose strong constraints on spatial pattern as well as on competitive relations and, thus, species coexistence. Therefore, the presence of clonal species within a plant community affects spatio-temporal dynamics and plant species diversity. We examined the distribution of plants with different clonal growth forms in the Northeast China Transect (NECT) and correlated plant species diversity with the importance of clonal plants, and the importance of phalanx and guerilla clonal plants. Phalanx clonal plants were more abundant in western communities where the altitude was higher and both the soil nitrogen contents and precipitation were relatively low. Whereas guerilla clonal plants were more abundant in the middle of the NECT where the precipitation, mean annual temperature and photosynthetically active radiation were relatively high. In the relatively productive temperate typical steppe, plant species diversity was negatively correlated with the importance of phalanx clonal plants and positively correlated with the importance of guerilla clonal plants. In relatively unproductive temperate desert steppe, plant species diversity was positively correlated with the importance of both phalanx and guerilla clonal plants.

Clonal plants and plant species diversity in wetland ecosystems in China

Abstract Clonal plants play important roles in maintaining wetland ecosystems in China. By analysing 108 wetland quadrats distributed throughout China, we evaluated (1) the importance of clonal growth forms in different Chinese wetlands, (2) how the abundance of clonal plants is related to climatic and geographical conditions, and (3) how plant species diversity is related to the abundance of clonal plants. Significant differences in clonal plant importance values were found between different regions of China. Clonal plants were more important in wetland ecosystems located towards the West and North and at higher elevations and, accordingly, experiencing a colder and drier climate. Plant species diversity showed a significant inverse correlation with the importance value of ‘guerilla’-type plants in most of the wetland regions. However, we found no significant correlation between plant species diversity and importance values of ‘phalanx’-type plants. In most Chinese wetlands, plant species diversity decreased with increasing importance of guerilla plants and also with an increase of the entire guild of clonal plants. In wetlands with low species richness, however, plant species diversity increased with increasing importance of guerilla plants and of all clonal plants together, suggesting that in these disturbed habitats clonal growth may facilitate the establishment of other, non-clonal wetland plants. Nomenclature: Anon. (1974); Anon. (1979–1999); Lang (1999).

Response of photosynthesis of different plant functional types to environmental changes along Northeast China Transect

Abstract Net photosynthesis (Pn), transpiration (E), stomatal conductance (gs), internal CO2 concentration (Ci), and water use efficiency (WUE) were examined on 215 species from eight plant functional types (PFTs) along a precipitation gradient in northeast China (the Northeast China Transect, or NECT). Among the eight PFTs, meadow steppe grasses had the highest rates of net photosynthesis and forest grasses the lowest and the following order of Pn was noted: meadow steppe grasses >typical steppe grasses >steppe shrubs >desert grasses >forest trees >forest shrubs >desert shrubs >forest grasses (P<0.05). Transpiration tended to be the highest in the steppe grasses and lowest in forest shrubs. Transpiration also decreased rapidly with the appearance of C3 desert species at the desert end. The forest tree PFT had lower Pn, E, gs than the steppe PFTs, whereas WUE values were somewhat greater in the forest tree PFT than the desert shrubs and grasses. Low Ci values along the steppe section (from 400 to 1100 km, east to west) indicated the presence of C4 species. Of all the PFTs, only shrubs and herbs were noted at all points along the transect. No clear relationship between Pn, E, gs, WUE of herb and shrub PFTs and annual precipitation was noted – low values were found at both the high and low precipitation ends of the transect. Highest values were noted when precipitation was intermediate.