Good mothers,bad mothers,and the nature of resistance to herbivory in <Emphasis Type="Italic">Solidago altissima</Emphasis>

Evidence of poor correspondence between an insect herbivore’s oviposition preferences and the performance of its offspring has generally been attributed either to maladaptive behavior of the insect mother or inadequate measurement by the researcher. In contrast, we hypothesize that many cases of “bad mothers” in herbivores may be a byproduct of the hierarchical way natural selection works on resistance in host plants. Epistatic selection on the components of resistance (i.e., antixenosis and antibiosis) may generate negative genetic correlations between the resistance components, which could counteract the efforts of herbivores to oviposit on the best hosts for the performance of their offspring. In common garden and greenhouse experiments, we measured aspects of antixenosis and antibiosis resistance in 26 genets of tall goldenrod, Solidago altissima, against two common herbivores: the gall-inducing fly Eurosta solidaginis and the spittlebug Philaenus spumarius. Goldenrod antixenosis and antibiosis were positively correlated against E. solidaginis and negatively correlated against P. spumarius. Analogously, population-wide preference–performance correlations were positive for the gall flies and negative for the spittlebugs. Several natural history differences between the two insects could make gall flies better mothers, including better synchrony of the phenologies of the flies and the host plant, the much narrower host range of the gall flies than the spittlebugs, and the more sedentary lifestyle of the gall fly larvae than the spittlebug nymphs. If these results are typical in nature, then negative genetic correlations in antixenosis and antibiosis in plants may often result in zero or negative population-wide correlations between preference and performance in herbivores, and thus may be an important reason why herbivorous insects often appear to be bad mothers.     

Temporal variation in herbivore host-plant preference and performance: constraints on host-plant adaptation

Temporal variation in the acceptability or suitability of plant genotypes to an herbivore has seldom been considered as a possible constraint limiting the adaptation of herbivores to particular plant genotypes, or the occurrence of a positive correlation between host-plant preference and offspring performance. In this study, we used data spanning 12 yr from the same 20 clones of goldenrod ( Solidago altissima ) to examine the temporal variation in oviposition preference and offspring performance of a stem-galling fly, Eurosta solidaginis . We found that the stem galler's preference for, or performance on, the different clones was uncorrelated between years of this study. Furthermore, we found that the relative rankings of clones changed by an average of 31% between successive years. We suggest that these consistently high year-to-year fluctuations in preference and performance by E. solidaginis are likely due to environmental factors (e.g., water and nutrient levels, or abundance of interspecific herbivores) that fluctuate over time and are known to differentially affect the acceptability and suitability of clones to herbivores; i.e., genotype×environment interactions. These results are significant because temporal fluctuations in host-plant preference and performance are likely to favor a more generalized diet by herbivorous insects.     

Evaluation of turfgrass species and cultivars for potential resistance to twolined spittlebug (Hemiptera: Cercopidae)

Potential resistance to the twolined spittlebug, Prosapia bicincta (Say), was evaluated among 56 turfgrass genotypes. Greenhouse, laboratory, and field bioassays identified differences in spittlebug survival and development, host preference and damage levels, and turfgrass tolerance to and ability to recover from pest induced injury. All centipede grasses demonstrated high levels of susceptibility, followed by bermudagrasses, seashore paspalums, and zoysiagrasses. Average nymphal survival to the adult stage ranged from 1.5 to 78.1%. Development required 38.1-62.0 d under greenhouse conditions, depending on plant taxa. Among seashore paspalums, nymphal survival to the adult stage was lowest and duration of development was longest on HI-1, 'Sea Isle 2000', 561-79, and 'Mauna Kea'. Reduced spittlebug survival and increased developmental times were also observed on the bermudagrasses BERPC 91-15 and 'Tifway'. Although zoysiagrasses supported spittlebug development and survival to the adult stage, developmental times were extended on the zoysiagrass cultivars 'Emerald' and 'El Toro'. Spittlebug preference varied with generation evaluated. First-generation spittlebugs inflicted the greatest damage on TC201 (centipede grass), 'Primavera' (bermudagrass), and 'Emerald' (zoysiagrass) in choice tests. In the fall, second-generation spittlebugs damaged TC201 (centipedegrass) and 'Sea Isle 1' (paspalum) most severely, whereas 561-79 (paspalum) and 'Emerald'(zoysiagrass) were less severely affected. Among taxa included in field trials, HI-1, 'Mauna Kea', 'Sea Isle 2000',and AP-14 paspalums, 'Tifway' bermudagrass, and 'Emerald' zoysiagrass were most tolerant (demonstrated the best regrowth potential following twolined spittlebug feeding).     

APHID DISTRIBUTION AND THE EVOLUTION OF GOLDENROD RESISTANCE

Although there is considerable evidence indicating that herbivory is detrimental to plant fitness, some recent studies of the evolution of plant resistance have concluded that insects do not impose selection on their host plants. A previously untested assumption that underlies most studies of the evolution of plant resistance is that insect distribution patterns are controlled directly by the effects of plant genotype on insect preference and performance. The experiments described here explicitly tested this assumption using the specialist herbivore Uroleucon tissoti (Homoptera: Aphididae) and its host plant Solidago altissima (Asteraceae). Measures of aphid preference and performance were used to predict aphid distribution patterns, and then the predicted distribution patterns were compared with the natural distribution pattern. Although goldenrod genotype had a strong effect on aphid distribution, aphid distribution was not controlled directly by the effect of goldenrod genotype on aphid preference and performance. Instead, a second experiment demonstrated that aphid and spittlebug (Philaenus spumarius and Lepyronia quadrangularis Homoptera: Cercopidae) distribution is controlled largely by genetic variation for resistance to a suite of “branch-causing” herbivores. These herbivores induce branching and aphids and spittlebugs are more abundant on branched plants than unbranched plants. These results indicate that any natural selection imposed by aphids and spittlebugs on goldenrod will depend on the presence or absence of branch-causing herbivores. Thus, selection for plant resistance may depend as much on the assemblage of insect species present as on the identity of each individual species.     

The influence of host plant variation and intraspecific competition on oviposition preference and offspring performance in the host races of Eurosta solidaginis

1. A series of experiments was conducted to measure the impact of plant genotype, plant growth rate, and intraspecific competition on the oviposition preference and offspring performance of the host races of Eurosta solidaginis (Diptera: Tephritidae), a fly that forms galls on Solidago altissima and Solidago gigantea (Asteraceae). Previous research has shown that both host races prefer to oviposit on their own host plant where survival is much higher than on the alternate host plant. In this study, neither host race showed any relationship between oviposition preference and offspring performance in choosing among plants of their natal host species. 2. The larval survival of both host races differed among plant genotypes when each host race oviposited on its natal host species. In one experiment, altissima host race females showed a preference among plant genotypes that was not correlated with offspring performance on those genotypes. In all other experiments, neither the altissima nor gigantea host race demonstrated a preference for specific host plant genotypes. 3. Eurosta solidaginis had a preference for ovipositing on rapidly growing ramets in all experiments, however larval survival was not correlated with ramet growth rate at the time of oviposition. 4. Eurosta solidaginis suffered high mortality from intraspecific competition in the early larval stage. There was little evidence, however, that females avoided ovipositing on ramets that had been attacked previously. This led to an aggregated distribution of eggs among ramets and strong intraspecific competition. 5. There was no interaction among plant genotype, plant growth rate, or intraspecific competition in determining oviposition preference or offspring performance.     

Negative impact of leaf gallers on arctic-alpine dwarf willow, Salix herbacea

According to available evidence, leaf gallers have only minor impacts on their plant hosts. We hypothesised that the relatively large leaf gallers formed by Eupontania sawflies on small, creeping arctic-alpine willows have a strong influence on their host plants. In this study, we specifically tested the effects of leaf galler (Eupontania aquilonis) on the survival and growth of dwarf willow (Salix herbacea) in a mountain snow-bed in northern Finland. We marked galled, leaves-removed and untreated ramets in experimental blocks. In the following year, we measured the growth and survival of the ramets. The mortality of galled shoots was approx. 40% higher, and the mortality of galled ramets approx. 25% higher than in the control ramets and in the leaf-removal treatment. The leaf biomass of galled ramets and the number of leaves were significantly less in galled ramets than in untreated or leaves-removed ramets. It is possible that galling causes fatal resource depletion of shoots in its host plant. The results show that leaf gallers are ecologically more influential than previously thought.     

Evolutionary impacts of the goldenrod ball gallmaker on Solidago altissima clones

Summary Three ramet clones of Solidago altissima were grown under greenhouse conditions to determine the effects of varying levels of attack by the goldenrod ball gallmaker (Eurosta solidaginis) on biomass allocation, leaf senescence rate and rhizome connections among ramets. The results, examined at both the individual ramet level and the clone level, showed that galled ramets became isolated from their clone through deterioration of rhizome connections. Gall effects were only observed at the ramet level although rhizome connection effects were detected at both the ramet and clone levels. The goldenrod ball gallmaker may therefore have little evolutionary impact on large clones but could appreciably affect newly established clones.     

Gall insects can avoid and alter indirect plant defenses

Parasitic species can dramatically alter host traits. Some of these parasite-induced changes can be considered adaptive manipulations that benefit the parasites. Gall-inducing insects are parasites well known for their ability to alter host-plant morphology and physiology, including the distribution of plant defensive compounds. Here it was investigated whether gall-inducing species alter indirect plant defenses, involving the release of volatile compounds that are attractive to foraging natural enemies. Using field and factorial laboratory experiments, volatile production by goldenrod (Solidago altissima) plants was examined in response to attack by two gall-inducing species, the tephritid fly Eurosta solidaginis and the gelechiid moth Gnorimoschema gallaesolidaginis, as well as the meadow spittlebug, Philaenus spumarius, and the generalist caterpillar Heliothis virescens. Heliothis virescens elicited strong indirect defensive responses from S. altissima, but the gall-inducing species and spittlebugs did not. More significantly, infestation by E. solidaginis appeared to suppress volatile responses to subsequent attack by the generalist caterpillar. The extensive control that E. solidaginis apparently exerts over host-plant defense responses may reduce the predation risk for the gall inducer and the subsequent herbivore, and could influence community-level dynamics, including the distribution of herbivorous insect species associated with S. altissima parasitized by E. solidaginis.     

Clonal integration affects growth and sediment properties of the first ramet generation,but not later ramet generations under severe light stress

克隆整合影响严重光胁迫下第一分株世代的生长和沉积物特征但不影响 后续分株世代的生长和沉积物特征 克隆整合通过缓冲环境压力和提高资源获取效率使克隆植物受益。然而,在一个克隆系统中,受益于克隆整合的连接分株世代的数量很少受到关注。我们进行了一个盆栽实验来评估沉水植物苦草 (Vallisneria natans)克隆系统内的生理整合程度,该克隆系统由一个母株和3个依次连接的后代分株组成。 母株生长在正常光照下,而后代分株被严重遮荫。母株与后代分株间的匍匐茎被切断或保持连接,但3个后代分株之间的连接仍然存在。与遮荫的后代分株连接时,苦草未遮荫的母株的光合能力显著增强,但其生物量积累大大减少。克隆整合显著增加了第一分株世代(相邻分株)的生物量积累和土壤的碳氮可用性、胞外酶活性和微生物生物量,但没有增加后续分株世代的这些特征。我们的结果表明,在严重光胁迫下,来自苦草母株的支持可能仅限于克隆系统中相邻的后代分株,这暗示着一个分株世代的效应。我们的结果有助于更好地理解克隆植物的层次结构和分段化。这些发现表明克隆整合程度在分株种群的生态相互作用中起着至关重要的作用。     

CLONAL INTEGRATION: NUTRIENT SHARING BETWEEN SISTER RAMETS OF SOLIDAGO ALTISSIMA (COMPOSITAE)

To test whether sharing of resources occurs among connected ramets of the tall goldenrod, Solidago altissima, we examined the extent of clonal integration for nutrients. In a greenhouse experiment, two-ramet clones were grown in a triad of connected pots so that nutrients could be supplied to either sister ramet or to their old rhizome (mother rhizome). Mother rhizomes and their associated roots shared nutrients with daughter ramets; however, any nutrient sharing that occurred between sister ramets was too little to significantly affect their growth. In addition, sister ramets not only competed for nutrients through parental connections, but larger ramets inhibited the growth of smaller ramets. We suggest that, for tall goldenrod, a clonal growth strategy in which nutrients are not shared among sister ramets may increase genet fitness by reducing the rhizome production of ramets in poor-nutrient microsites. Consequently, the genet would produce relatively fewer ramets in unfertile areas and make better use of heterogeneous nutrient resources.     

Costs and benefits of gregarious feeding in the meadow spittlebug, Philaenus spumarius

Abstract.  1. To study the function of group living in a non-social animal, costs and benefits of feeding in different group sizes (one to eight) were measured for nymphs of the meadow spittlebug ( Philaenus spumarius ) on tall goldenrod ( Solidago altissima ).
2. In a controlled greenhouse experiment, the number of nymphs per plant had a significant influence on their mass as adults. Adult mass increased as group size went from one to five nymphs per plant, but fell as the group size increased beyond five. In contrast, the density of nymphs on a plant had no effect on the duration of nymphal development.
3. Nymphal mortality increased linearly as the number of nymphs increased from one to eight per plant.
4. These performance results suggest that the optimal group size of spittlebug nymphs reflects a compromise between bottom-up influences (e.g. overcoming physical barriers to feeding on xylem) and within-trophic-level influences (e.g. avoiding competition for limited plant resources).
5. In a survey of local populations of goldenrod, spittlebug nymphs were found to be slightly but significantly aggregated among host-plant individuals. In addition, when more than one nymph occurred on a plant, the nymphs tended to feed in shared spittle masses.
6. It is concluded that the aggregated dispersion pattern of spittlebug nymphs reflects an adaptive balance of coping with potentially conflicting demands from different trophic levels.     

Integration in the clonal plant <Emphasis Type="Italic">Eriophorum angustifolium</Emphasis>: an experiment with a three-member-clonal system in a patchy environment

A clonal plant in heterogeneous environments is usually expected to profit from resource exchange via a clonal network where ramets placed in contrasting environments can specialise so to acquire the most abundant resources. An experiment was designed using the three member clonal system of Eriophorum angustifolium, which consisted of one parent ramet growing in a resource poor environment and two offspring: one was limited in growth by nutrients while the other was light limited; the contrast in availability of limited resources between the offspring ramets was high, medium or none, with the system either connected or severed. The total resource availability was the same in all treatments. We proposed four possible scenarios for the system: offspring ramets will share resources via the deficient parent ramet, and the whole clone will profit from the contrasting environment (scenario 1); offspring ramets will support exclusively the parent ramet, and the whole clone will profit from a homogeneous environment (scenario 2); offspring ramets will stop the export of the limiting resource to the parent ramet, with split and connected treatments not differing (scenario 3); and offspring ramets will exhaust the carbon stored in the biomass of the parental ramet; offspring ramet will profit from connection (scenario 4). In the experiment, the limiting resources were sent to the strongest sink (scenario 2). The parent ramet growing in a deficient environment received the highest support in the treatment where both offspring ramets were growing in the same conditions (no-contrast treatment). Production of new shoots, but not biomass of whole clone, was supported in a homogenous environment. The experiment revealed that multiple stresses might prohibit free exchange of limiting resources via the clonal network and supports the idea that experimental studies on more complex clones are essential for understanding the costs and benefits of clonal growth.     

Quantitative genetics of preference and performance on chickpeas in the noctuid moth, Helicoverpa armigera

If a novel, resistant host-plant genotype arises in the environment, insect populations utilising that host must be able to overcome that resistance in order that they can maintain their ability to feed on that host. The ability to evolve resistance to host-plant defences depends upon additive genetic variation in larval performance and adult host-choice preference. To investigate the potential of a generalist herbivore to respond to a novel resistant host, we estimated the heritability of larval performance in the noctuid moth, Helicoverpa armigera, on a resistant and a susceptible variety of the chickpea, Cicer arietinum, at two different life stages. Heritability estimates were higher for neonates than for third-instar larvae, suggesting that their ability to establish on plants could be key to the evolution of resistance in this species; however, further information regarding the nature of selection in the field would be required to confirm this prediction. There was no genetic correlation between larval performance and oviposition preference, indicating that female moths do not choose the most suitable plant for their offspring. We also found significant genotype by environment interactions for neonates (but not third-instar larvae), suggesting that the larval response to different plant genotypes is stage-specific in this species.     

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.     

Aphid and ladybird beetle abundance depend on the interaction of spatial effects and genotypic diversity

Intraspecific variation and genotypic diversity of host-plants can affect the structure of associated arthropod communities and the dynamics of populations. Similarly, neighboring plants can also affect interactions between host-plants and their associated arthropods. However, most studies on the effects of host-plant genotypes have largely ignored the potential effects of neighboring host-plants on arthropod communities. In this study, we used a common garden experiment to ask how spatial effects of neighboring patches, along with genotype identity and genotypic diversity in tall goldenrod (Solidago altissima), affect the abundances of a common goldenrod herbivore (Uroleucon nigrotuberculatum) and their dominant predator (Harmonia axyridis, a ladybird beetle). Aphid abundance varied 80-fold among genotypes, while ladybird beetle abundance was not affected by genotype identity. Additionally, there were strong effects of neighboring plots: aphid abundance in a focal plot was positively correlated to aphid abundance in nearby plots, suggesting strong spatial patterning in the abundance of aphids. Neither aphid nor ladybird beetle abundance was affected by genotypic diversity. However, focal plot genotypic diversity mediated the strength of the neighborhood effect (i.e., strong effects for genotype polyculture focal plots and weak effects for genotype monoculture focal plots). Our results show that aphids were directly influenced by host-plant genotype identity while ladybird beetles responded mainly to prey abundance, and suggest that genotypic diversity can influence the effects of spatial processes on the plant-herbivore interactions.     

Dynamics of distribution and performance of ramets constructing genets: a demographic�Cgenetic study in a clonal plant, Convallaria keiskei

Background and Aims

In clonal plants producing vegetative offspring, performance at the genet level as well as at the ramet level should be investigated in order to understand the entire picture of the population dynamics and the life history characteristics. In this study, demography, including reproduction and survival, the growth patterns and the spatial distributions of ramets within genets of the clonal herb Convallaria keiskei were explored.

Methods

Vegetative growth, flowering and survival of shoots whose genets were identified using microsatellite markers were monitored in four study plots for 3 years (2003–2005). The size structures of ramets in genets and their temporal shifts were then analysed. Their spatial distributions were also examined.

Key Results

During the census, 274 and 149 ramets were mapped in two 1 × 2 m plots, and 83 and 94 ramets in two 2 × 2 m quadrats. Thirty-eight genotypes were identified from 580 samples. Each plot included 5–18 genets, and most ramets belonged to the predominant genet(s) in each plot. Shoots foliated yearly for several years, but flowering ramets did not have an inflorescence the next year. A considerable number of new clonal offspring persistently appeared, forming a bell-shaped curve of the size structure of ramets in each genet. Comparing the structures modelled by the normal distributions suggested variation among ramets belonging to a single genet and variation among genets. Furthermore, spatial analyses revealed clumped and distant distributions of ramet pairs in a genet, in which the distant patterns corresponded to the linearly elongating clonal growth pattern of this species.

Conclusion

Characteristics of ramet performances such as flowering and recruitment of clonal offspring, in addition to growth, played a large part in the regulation of genet dynamics and distribution, which were different among the studied genets. These might be characteristics particularly relevant to clonal life histories.Key words: Clonal plant, Convallaria keiskei, demography, genet, genetic identification, growth pattern, life history, ramet, spatial distribution     

Associative nitrogen fixation, C4 photosynthesis, and the evolution of spittlebugs (Hemiptera: Cercopidae) as major pests of neotropical sugarcane and forage grasses

Neotropical grass-feeding spittlebugs of several genera are important pests of pasture grasses from the southeastern USA to northern Argentina, and of sugarcane from southern Mexico to southern Brazil, causing estimated reductions of up to 70% in yield and estimated monetary losses of 840-2100 million US dollars annually. With few exceptions, the species badly damaged by these spittlebugs are introduced C4 grasses that exhibit associative nitrogen fixation. This study synthesizes evidence that the pest status of many tropical and subtropical grass-feeding spittlebugs is linked to associative N-fixation in their C4 hosts. Recognition that associative N-fixation is a major factor in spittlebug host preferences should deepen understanding of spittlebug agricultural ecology and facilitate efforts to combat spittlebug pests. In particular, spittlebugs should be susceptible to manipulation of xylem transport solutes. However, reduction of nitrate fertilizer rates, increase in ammonium fertilizer rates, or enhancement of associative N-fixation as a consequence of genetic engineering could make hosts more susceptible to spittlebug attack. Because of their predilection for C4 grasses, spittlebugs present a clear counterexample to the hypothesis that herbivores prefer C3 plants to C4 plants. Finally, it appears that declines in atmospheric carbon dioxide levels during recent geological history promoted the proliferation of C4 grasses. This, compounded by human agricultural activities, has driven an ecological and evolutionary radiation of grass-feeding spittlebugs that presents continuing opportunities for the evolution of spittlebug pests.     

Differences in the structure, growth and survival of Parasenecio yatabei ramets with contrasting water relations on the slope of a stream bank

Parasenecio yatabei (Asteraceae), a summer-green perennial herb, is widely distributed on sloping mountain stream banks in cool-temperate zone forests of Japan. We investigated the growth pattern, leaf longevity and leaf water relations of vegetatively independent plants (ramets) growing in two contrasting soil water conditions, that is, upper and lower stream banks (U ramets and L ramets, respectively). The objective of the present study was to clarify the physiological and morphological responses of the ramets to soil water conditions. Dry matter allocation to subterranean parts was higher in U ramets than in L ramets. The U ramet leaves survived for approximately 2 months longer than L ramet leaves. The ratio of subterranean part to aerial part dry matter was greater in U ramets than L ramets. Leaf mass per leaf area (LMA) tended to be greater in U ramets than L ramets throughout the growing season. The leaf bulk modulus of elasticity at full hydration was significantly higher in U ramets. Thus, ramet growth patterns and morphological traits varied with changing soil water conditions. The greater longevity of U ramet leaves may play a role in compensating for the reduced annual net carbon gain caused by lower photosynthetic activity. U ramets growing in environments with less water availability achieved high water-use efficiency by a high passive water absorption capacity via a progressed root system and high productivity via longer leaf longevity with higher LMA and elasticity. Therefore, P. yatabei growing along mountain streams could have the ability to colonize the upper bank through higher survivorship based on these traits.     

CLONE STRUCTURE IN FOUR SOLIDAGO ALTISSIMA (ASTERACEAE) POPULATIONS: RHIZOME CONNECTIONS WITHIN GENOTYPES

We studied clone structure and degree of genotypic mixing of Solidago altissima L. (Asteraceae) clones in four old fields near Ithaca, New York. The fields differed in time from agricultural abandonment and were approximately 1, 5, 20, and 35 years old. In the three older fields, three 0.75 m² plots were excavated intact and rhizome connections among ramets were mapped. In the youngest field 30 ramets were dug up singly. The genotype of all ramets was determined using electrophoresis of four polymorphic enzyme systems. Fields differed in the number and dispersion of genotypes within plots, and the degree of connection among ramets in the same clone. The one-year-old field was composed of single ramet genotypes which had probably established from seed the previous year. The five-year-old field contained many small contiguous clones of S. altissima with highly interconnected ramets. In the oldest two fields clones were highly intermixed and ramets of the same genotype were not extensively interconnected. These results demonstrate that clones of S. altissima display considerable phenotypic variability between fields and patterns of clone development may differ. The causes of this variability remain to be identified. We suggest that either selection for different genotypes or changing habitat conditions during succession may lead to changes in clone form.     

Exploitation of patchy soil water resources by the clonal vine <Emphasis Type="Italic">Ficus tikoua</Emphasis> in karst habitats of southwestern China

The karst habitats of southwestern China are characterized by a highly heterogeneous distribution of water resources. We hypothesized that the clonal integration between connected ramets of the clonal vine Ficus tikoua was an important adaptive strategy to the patchy distribution of water resources in these habitats. We grew ramet pairs (each consisting of a parent and an offspring ramet) in both homogeneously and heterogeneously watered conditions. The offspring ramets were well-watered, whereas their connected parent ramets were randomly assigned to four water treatments: well-watered, mild water stress, moderate water stress, and severe water stress. Increasing water stress decreased leaf water potential, relative water content, net assimilation rate, maximum quantum yield of PSII (F _v/F _m), and biomass of the parent ramets. Subjecting the parents to water stress significantly increased root biomass and root mass ratio (RMR) of their offspring ramets. Exploitation of plentiful water resources through the increased adventitious roots connected to another soil patch permitted the complete restoration of water relations and photosynthetic capacity of offspring ramets after an initial depression. Water relations and gas exchange of the parents were not affected by the water supply to their connected offspring ramets, suggesting that offspring ramets hardly exported water to the stressed parents. However, net assimilation rate and proline content of the offspring ramets increased when they were connected to water-stressed parents. The compensatory photosynthetic responses of offspring ramets connected to stressed parents revealed an increasing trend as the experiment progressed. Morphological and physiological plasticity of F. tikoua in response to heterogeneous water resources allow them to adapt to karst habitats and be suitable candidates for vegetation restoration projects.