Is a new invasive herb emerging? Molecular confirmation and preliminary evaluation of natural hybridization between the invasive Sphagneticola trilobata (Asteraceae) and its native congener S. calendulacea in South China

Sphagneticola trilobata is an invasive herb in South China. Here, we reported some morphological intermediates between S. trilobata and the indigenous congener S. calendulacea in the field for the first time, which were suspected to be hybrids. Morphological and molecular methods were used to test this hybrid origin hypothesis, and eco-physiological parameters were measured to evaluate the hybrid??s invasion potential. The hybrid origin hypothesis was confirmed by analysis of microsatellite and nrITS sequences. Bayesian assignment analysis based on six microsatellite loci genotypes separated the studied individuals into three groups, which corresponded to the two parent species and the putative hybrids that were identified on the basis of leaf morphology. The hybrid individuals were categorized as F1 hybrids with membership coefficients (Q_i values) close to 0.5. Phylogenetic analysis based on nrITS data showed that the putative hybrid individuals possessed two types of species-level divergent clones, and were inseparate with S. trilobata clade and S. calendulacea clade, respectively. The maternally-inherited chloroplast marker atpB-rbcL demonstrated that the invasive species functioned as the pollen donors. Ecophysiological studies on photosynthesis, specific leaf area (SLA), leaf nutrient concentration, and construction cost in the parental species and the hybrid demonstrated that the hybrid showed slightly lower or equivalent growth potential compared to the invasive species and significant higher energy-use efficiency than both parents. As such, it seems that a new invasive herb is emerging. As a consequence of this hybridization, the native species S. calendulacea may be disappearing rapidly, which presents a major concern for conservation efforts.     

遮荫对南美蟛蜞菊、蟛蜞菊及其自然杂交种叶片显微结构及叶绿体超微结构的影响

利用JSM-6360LV型扫描电镜和JEM-1010型透射电镜,观察了南美蟛蜞菊、蟛蜞菊及其自然杂交种新近成熟和老熟叶片的解剖结构及叶绿体超微结构。结果表明:遮荫后该杂交种与其亲本新近成熟叶片均表现为上下表皮气孔密度、叶片总厚度及上下表皮厚度、栅栏组织、海绵组织厚度减小,叶绿体肿胀变形,基粒片层垛叠程度增加,淀粉粒增多变大;遮荫后杂交种老熟叶片总厚度及上表皮、栅栏组织、海绵组织厚度增加,入侵种的下表皮厚度及本地种的上表皮厚度增加,叶绿体超微结构在遮荫后均出现严重损伤,基粒片层类囊体结构边缘溶解等。说明三种蟛蜞菊属物种及各物种不同叶龄叶片对弱光条件的响应存在差异;杂交种叶片显微及超微结构在不同光照下的变化介于亲本之间,对遮荫环境能较好适应。     

异质光照环境下克隆整合对南美蟛蜞菊和蟛蜞菊种间关系的影响

很多入侵植物具有克隆性,克隆整合对入侵克隆植物生长和繁殖具有重要的贡献。自然界中,植物生长和繁殖所需的各种资源如光照、水分和矿质养分等在空间上分布通常是异质的,但关于异质环境下克隆整合对入侵植物和本土同属植物种间关系影响的研究相对缺乏。通过温室控制实验,将入侵植物南美蟛蜞菊(Wedelia trilobata)和同属本土植物蟛蜞菊(W.chinensis)的分株对单独种植或者混合种植在异质性光照条件下,同时通过保持或者切断分株之间的连接来控制克隆整合效应的有无,研究异质光照环境下克隆整合对南美蟛蜞菊和蟛蜞菊种间关系的影响。克隆整合对南美蟛蜞菊和蟛蜞菊的生长和繁殖都有促进作用,且南美蟛蜞菊比蟛蜞菊从克隆整合中获益更多。与单独种植相比,两者混种对南美蟛蜞菊的叶生物量有显著影响,而对本地种蟛蜞菊的根生物量有显著影响。克隆整合和种间关系对南美蟛蜞菊的总生物量和叶生物量产生了显著的交互作用,而对蟛蜞菊各指标无显著影响。克隆整合状态显著影响了南美蟛蜞菊和蟛蜞菊的种间关系。这些结果表明,异质环境下克隆整合可以改变入侵植物南美蟛蜞菊和本土植物蟛蜞菊的生长性状及种间关系。     

A hybrid of the invasive plant Sphagneticola trilobata has similar competitive ability but different response to nitrogen deposition compared to parent

Hybridization between introduced and related native species has been suggested to be a key driving force of evolutionary processes in invasions that may be accelerated by increasing nitrogen (N) deposition. We carried out two experiments to compare the competitive ability of a newly-reported putative hybrid (Sphagneticola calendulacea × Sphagneticola trilobata) to that of its invasive paternal parent and investigate its response to different N additions. Growth of the hybrid increased by 39.62 ± 15.69 % when grown with the native parent but showed no significant changes when grown with the invasive parent. Growth of invasive S. trilobata showed no significant changes when grown with the other two taxa. The hybrid and invasive parent inhibited the growth of the native parent by 33.27 ± 15.26 % and 78.56 ± 3.13 %, respectively. Low level N addition (50 kg N ha^?1) enhanced the growth of both the hybrid and native parent, whereas medium and high levels of N additions (150, 300 kg N ha^?1) had no effects on hybrid growth but decreased native parent growth. However, all N treatments had no significant effects on the growth of the invasive parent. Our results suggest that the hybrid is an equal competitor to its invasive parent and is more susceptible to N deposition. Therefore, this hybrid could be a threat to native diversity but may decline more than the invasive parent under high N conditions.     

Altering Light Availability to Restore Invaded Forest: The Predictive Role of Plant Traits

Many studies have demonstrated that reduced light availability, which can be manipulated at local scales by planting or seeding canopy species, can curtail the growth of invasive species and promote the growth of native species. Species differences in functional traits, such as light use and stress tolerance, may be used to determine how native and invasive species will respond to these resource manipulations. We altered light availability in a mesic Hawaiian forest understory and found that low light levels reduced the biomass and growth of two invasive grasses (Pennisetum clandestinum and Ehrharta stipoides) relative to two native shrubs (Pipturus albidus and Coprosma rhynchocarpa) and two native canopy species (Metrosideros polymorpha and Acacia koa). Native species generally displayed traits associated with shade tolerance (high quantum yield, chlorophyll content, and leaf mass per area), whereas the two invasive grasses displayed traits associated with shade intolerance (high photosynthetic rate and growth rate). Several key traits pertaining to light acquisition and shade tolerance (quantum yield, chlorophyll content, and leaf mass per area) predicted seedling survival in low‐light treatments. Our data suggest that differences in light use among native and invasive species can help to determine the utility of resource manipulation as a restoration tool and, more specifically, to predict which native species will be optimal for restoration efforts that manipulate light availability.     

Plasticity in seedling morphology,biomass allocation and physiology among ten temperate tree species in response to shade is related to shade tolerance and not leaf habit

• Mechanisms of shade tolerance in tree seedlings, and thus growth in shade, may differ by leaf habit and vary with ontogeny following seed germination. To examine early responses of seedlings to shade in relation to morphological, physiological and biomass allocation traits, we compared seedlings of 10 temperate species, varying in their leaf habit (broadleaved versus needle‐leaved) and observed tolerance to shade, when growing in two contrasting light treatments – open (about 20% of full sunlight) and shade (about 5% of full sunlight).
• We analyzed biomass allocation and its response to shade using allometric relationships. We also measured leaf gas exchange rates and leaf N in the two light treatments.
• Compared to the open treatment, shading significantly increased traits typically associated with high relative growth rate (RGR) – leaf area ratio (LAR), specific leaf area (SLA), and allocation of biomass into leaves, and reduced seedling mass and allocation to roots, and net assimilation rate (NAR). Interestingly, RGR was not affected by light treatment, likely because of morphological and physiological adjustments in shaded plants that offset reductions of in situ net assimilation of carbon in shade. Leaf area‐based rates of light‐saturated leaf gas exchange differed among species groups, but not between light treatments, as leaf N concentration increased in concert with increased SLA in shade.
• We found little evidence to support the hypothesis of a increased plasticity of broadleaved species compared to needle‐leaved conifers in response to shade. However, an expectation of higher plasticity in shade‐intolerant species than in shade‐tolerant ones, and in leaf and plant morphology than in biomass allocation was supported across species of contrasting leaf habit.

     

Fluctuated water depth with high nutrient concentrations promote the invasiveness of Wedelia trilobata in Wetland

The distribution of invasive and native species in wetlands is determined by hydrological conditions; whereas conditions such as water depth fluctuations, variations in the nutrient concentrations are expected to affect the growth and physiological traits of plants. For the assessment of such effects, we conduct greenhouse experiment with three factors; 1) water depth of 5 cm and 15 cm (static and fluctuated); 2) three levels of nutrient concentrations (i) full‐strength Hoagland solution (N1), (ii) ¼‐strength Hoagland solution (N2), and (iii) ¹/₈‐strength Hoagland solution (N3); and 3) species, invasive Wedelia trilobata (L.) and its congener, native Wedelia chinensis (Osbeck.) under mono and mixed culture. Water depth of 5 cm combined with any of the nutrient treatments significantly restrained the photosynthesis, intracellular CO₂ concentration and leaf chlorophyll of both W. trilobata and W. chinensis. Increase in the water depth to 15 cm with low‐nutrient treatment N3 did not sustain the physiological traits of W. chinensis under mono and mixed planting. A great loss was noted in the growth of W. chinensis at 15 cm static and fluctuated water depth with low‐nutrient treatment (N3) and under mixed culture. In addition, water depth fluctuations with both low‐ and high‐nutrient treatments significantly affected the root‐shoot ratio, relative growth rate, and interspecific interaction among these two species. W. trilobata benefited more from competitive interaction index (CII) under fluctuated water depth at 15 cm with high nutrients, and the value of CII was clearly positive. Therefore, higher competitive ability may contribute to the invasiveness of W. trilobata in wetlands.     

Elevated CO<Subscript>2</Subscript> increases energy-use efficiency of invasive <Emphasis Type="Italic">Wedelia trilobata</Emphasis> over its indigenous congener

Increasing atmospheric CO₂ concentration is regarded as an important factor facilitating plants invasions by stimulating invasive species growth. However, the physiological mechanisms by which invasive plants increase at the expense of existing native plants are poorly understood. Plant growth is always related to energy-use process including energy assimilation and expenditure, and thus examination of energetic properties could provide mechanistic insight into growth responses to increased CO₂. The aims of this study were to examine the effect of rising CO₂ on the growth and energetic properties of alien invasive species (Wedelia trilobata (L.) Hitchc.) and its native congener (Wedelia chinensis (Osbeck.) Merr.) in South China, and to determine if the specific energetic properties of invasive species at elevated CO₂ favoring its growth. Elevated CO₂ stimulated a greater increase in biomass production for invasive W. trilobata (58.9%) than for its indigenous congener (48.1%). Meanwhile, elevated CO₂ altered the energetic properties differently upon species. For invasive W. trilobata, elevated CO₂ significantly increased total energetic gain via photosynthetic activity (A _total), but decreased energetic cost of biomass construction (CC), and thus enhanced photosynthetic energy-use efficiency (PEUE) by 85.3%. In contrast, the indigenous W. chinensis showed a slight increase in PEUE by 43.8%. Additionally, W. trilobata individuals grown in elevated CO₂ increased energy allocation towards stems. Statistic analysis revealed significant associations between growth characteristics (relative growth rate and biomass) and energetic properties (CC and PEUE), suggesting the greater growth stimulation in invasive species could be partly explained by its specific energetic properties in elevated CO₂ concentration. The invasive species showed a greater increase in energy-use efficiency under elevated CO₂, which consequently facilitated its growth. It might be a physiological mechanism promoting success of invasion with ongoing increase in atmospheric CO₂ concentration.     

Silk filaments enhance the settlement of stream insect larvae

The recombination that follows natural hybridization may produce hybrid genotypes with traits that are intermediate or extreme relative to the parental species, and these traits may influence the relative fitness of the hybrids. Here we examined leaf ecophysiological traits that may influence fitness patterns in a natural plant hybrid zone. We compared the biochemical photosynthetic capacity of Ipomopsis aggregata, I. tenuituba, and early generation hybrids, as well as their photosynthetic responses to varying light and temperature, two abiotic factors found to differ among sites along the hybrid zone. In general, ecophysiological traits expressed in these plants were consistent with their natural habitat, even when grown under common greenhouse conditions. I. tenuituba reached higher photosynthetic rates (A) at higher light levels than I. aggregata, and also had a higher optimal temperature for photosynthesis (T _opt). This suite of traits may reflect adaptations to the more exposed, rocky sites where I. tenuituba is found, compared to the more vegetated, mesic I. aggregata site. Hybrids had characters that were largely intermediate or tenuituba-like, but particular individual hybrids were extreme for some traits, including light saturation level, light-saturated A, and T _opt. Many of these traits are consistent with adaptations reported for plants found in warm, dry sites, so they may put certain hybrids at an advantage at the relatively xeric center of the natural hybrid zone.     

Vigour-related traits and immunity in hybrids of evolutionary divergent cyprinoid species: advantages of hybrid heterosis?

Hybrid advantage, described as the superiority of hybrids in some traits over their parents and termed the “heterosis effect,” is widely documented in the case of reciprocal crosses of parental species (i.e., hybrids representing the F1 generation). In fish, high survival, fast growth and better health status have been widely documented in F1 hybrids. Nonetheless, the effects of interspecific hybridization on vigour, physiology and immunity-related traits in fish are largely unknown, especially concerning native systems of coexisting parental and hybrid genomes in the same habitat. The present study examined the potential physiological and immune aspects of hybrid heterosis by comparing condition status (measured especially by indexes), haematological profile, glucose concentration and selected parameters of non-specific and specific immunity between the evolutionarily divergent non-congeneric cyprinoid species Abramis brama and Rutilus rutilus and their hybrids representing the F1 generation, all of them caught in nature. Clear differences were documented for vigour-related, physiological and immune parameters between the two divergent species. Hybrids generally tended to express intermediate characters of the measured traits, likely generated by the evolutionary divergence of the hybridizing species; nonetheless, for some traits, hybrids exhibited a character that was more similar to one parental species than to the other. This was interpreted as the heterozygote advantage for F1 hybrids. It is suggested that a maternally inherited genetic background may potentially influence the expression of some branches of non-specific immunity or other aspects related to the fish health status.     

The production of hybrids with high ecological amplitude between exotic Spartina densiflora and native S. maritima in the Iberian Peninsula

Aim Hybridization is a common and potent mechanism of plant evolution that has the potential to be evolutionary significant in its own right, and hybrids are common between invasive and native congeneric species. Our aims were to document the existence and nature of new Spartina hybrids arising between introduced Spartina densiflora and native S. maritima in tidal marshes of the Iberian Peninsula, to examine the actual and potential range of hybrids in the intertidal zone, and to analyse the seed set of hybrids. Location South‐West Iberian Peninsula. Methods Hybrids were characterized using chloroplast and nuclear DNA, and ploidy assessments. The ecological tolerance of the hybrids was studied using vegetation surveys and transplant experiments. Results We found that both parental species have been seed parents to hybrids and that all hybrids had an additive pattern of species‐specific nuclear markers consistent with F₁ hybrids. Hybrid chromosome numbers varied between 2n = ca. 65 and 2n = ca. 94–97, while S. maritima had 2n = ca. 60 and S. densiflora had 2n = ca. 70. Hybrids grew in three discrete locations along the intertidal zone but were capable of growing throughout the ranges of both parental species in transplanted gardens, and in most cases, grew better than the parental species. While the potential exists for the origination of another invasive Spartina hybrid species, thus far hybrid plants are not fertile, limiting their ability to invade and spread. Main conclusions We recommend the eradication of all of the currently quite limited hybrid cordgrass and to fight the S. densiflora invasion in the Iberian Peninsula to prevent the origination of a new invasive allopolyploid Spartina species.     

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

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

Needle morphological evidence of the homoploid hybrid origin of Pinus densata based on analysis of artificial hybrids and the putative parents,Pinus tabuliformis and Pinus yunnanensis

Genetic analyses indicate that Pinus densata is a natural homoploid hybrid originating from Pinus tabuliformis and Pinus yunnanensis. Needle morphological and anatomical features show relative species stability and can be used to identify coniferous species. Comparative analyses of these needle characteristics and phenotypic differences between the artificial hybrids, P. densata, and parental species can be used to determine the genetic and phenotypic evolutionary consequences of natural hybridization. Twelve artificial hybrid families, the two parental species, and P. densata were seeded in a high‐altitude habitat in Linzhi, Tibet. The needles of artificial hybrids and the three pine species were collected, and 24 needle morphological and anatomical traits were analyzed. Based on these results, variations in 10 needle traits among artificial hybrid families and 22 traits among species and artificial hybrids were predicted and found to be under moderate genetic control. Nineteen needle traits in artificial hybrids were similar to those in P. densata and between the two parental species, P. tabuliformis and P. yunnanensis. The ratio of plants with three needle clusters in artificial hybrids was 22.92%, which was very similar to P. densata. The eight needle traits (needle length, the mean number of stomata in sections 2 mm in length of the convex and flat sides of the needle, mean stomatal density, mesophyll/vascular bundle area ratio, mesophyll/resin canal area ratio, mesophyll/(resin canals and vascular bundles) area ratio, vascular bundle/resin canal area ratio) relative to physiological adaptability were similar to the artificial hybrids and P. densata. The similar needle features between the artificial hybrids and P. densata could be used to verify the homoploid hybrid origin of P. densata and helps to better understand of the hybridization roles in adaptation and speciation in plants.     

遮阴对骆驼刺叶性状和水分生理的影响

以塔克拉玛干沙漠南缘荒漠绿洲过渡带优势种骆驼刺为试验材料,研究正常光照(NL)、中度遮阴(MS)和重度遮阴(SS)3种不同遮阴环境对骆驼刺(Alhagi sparsifoliashap.)叶厚、单叶面积、比叶面积、比叶质量等叶性状参数和相对含水量、水势、失水速率、气孔导度、蒸腾速率以及水分利用效率等水分生理参数的影响。结果显示:随着光照强度的降低,骆驼刺的叶片厚度(LT)、比叶质量(LMA)、净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)和水分利用效率(WUE)明显降低,而单叶面积(LA)、比叶面积(SLA)、相对含水量(RWC)、失水速率(RWL)和水势(WP)明显升高;各叶性状参数与水分生理参数之间均有极显著的相关性。说明:骆驼刺在遮阴环境下,其叶性状特征主要通过单叶面积和单叶干物质积累的变化来响应遮阴环境;其水分生理特征主要是牺牲对水分胁迫具有较好抗逆性的水分生理特征并通过气孔调节和更多的水分消耗用于维持一定的光合能力来响应遮阴环境。叶性状与水分生理参数相关性分析表明,遮阴环境下骆驼刺叶性状变化主要通过影响RWL、WP、Tr和WUE进而影响其水分生理特征的变化。因此,建议可利用遮阴措施对极端干旱区荒漠植物骆驼刺进行植被恢复,但其遮光度应设置在70%—80%自然光之间。     

Shade tolerance and herbivory are associated with RGR of tree species via different functional traits

• Relative growth rate (RGR) plays an important role in plant adaptation to the light environment through the growth potential/survival trade‐off. RGR is a complex trait with physiological and biomass allocation components. It has been argued that herbivory may influence the evolution of plant strategies to cope with the light environment, but little is known about the relation between susceptibility to herbivores and growth‐related functional traits.
• Here, we examined in 11 evergreen tree species from a temperate rainforest the association between growth‐related functional traits and (i) species’ shade‐tolerance, and (ii) herbivory rate in the field. We aimed at elucidating the differential linkage of shade and herbivory with RGR via growth‐related functional traits.
• We found that RGR was associated negatively with shade‐tolerance and positively with herbivory rate. However, herbivory rate and shade‐tolerance were not significantly related. RGR was determined mainly by photosynthetic rate (A_max) and specific leaf area (SLA). Results suggest that shade tolerance and herbivore resistance do not covary with the same functional traits. Whereas shade‐tolerance was strongly related to A_max and to a lesser extent to leaf mass ratio (LMR) and dark respiration (R_d), herbivory rate was closely related to allocation traits (SLA and LMR) and slightly associated with protein content.
• The effects of low light on RGR would be mediated by A_max, while the effects of herbivory on RGR would be mediated by SLA. Our findings suggest that shade and herbivores may differentially contribute to shape RGR of tree species through their effects on different resource‐uptake functional traits.

     

Morphological plasticity and ecophysiological response of ground ivy (Glechoma hederacea,Lamiaceae) in contrasting natural habitats within its native range

Abstract

Some species intrinsically have a high invasiveness capacity, shown by high phenotypic plasticity and rapid growth, enabling a wide distribution across their native habitats and successful invasion in the introduced range. For such species, information from native habitats is critically important. An example is Glechoma hederacea, native to Eurasia but introduced and widespread in the USA. Our main objective was to investigate variation in traits of native G. hederacea populations across contrasting habitats: open, forest edge and understory. Vegetation was sampled and the ecophysiological and morphological traits were measured with accompanying environmental parameters. Results showed that in native habitats environmental conditions cover wide gradients of light and soil moisture. Plants had the highest cover in nutrient-rich, shaded habitats, representing the optimal habitat, indicating shade tolerance of G. hederacea. Plants from forest understory exhibited strong similarities in investigated traits to plants from the forest edge, even though this was a drier, sunnier habitat. Plants from open, sunny habitats experienced stress as indicated by the quantum efficiency of PSII and significantly higher sexual reproduction. Results show that G. hederacea is moderately tolerant simultaneously to shade and drought, a characteristic that has been reported for numerous invasive species, while at the same time it shares some characteristics with weedy plants.     

SHADE TOLERANCE AND ITS EFFECT ON THE SEGREGATION OF TWO SPECIES OF LOUISIANA IRIS AND THEIR HYBRIDS

Iris fulva Ker. Gawler and Iris hexagona Walter have overlapping geographic ranges in Louisiana. In areas of overlap hybrids are fairly common. Iris hexagona occupies the borders of freshwater marshes of southern Louisiana while I. fulva can be found farther north along edges of natural levees, canals and swamps. Where the natural levee penetrates the marsh, natural hybridization can occur between I. hexagona and I. fulva. It has been suggested that one principal explanation for the segregation of the two species is that I. fulva grows best in semishade and I. hexagona grows best in full sun. A greenhouse study was conducted using rhizomes collected from the field to test this hypothesis and determine the relative shade tolerance of two natural hybrid types. Iris fulva, I. hexagona, and the two hybrid taxa were grown under 0% (control), 50% (medium shade), and 80% (high shade) reduction of sunlight for 6 months and then harvested. Iris fulva was found to be more tolerant of shading than I. hexagona and the two hybrids. Further, I. fulva was found to grow as well in control as in medium shade. Both hybrid taxa were more shade tolerant than I. hexagona. Iris hexagona was greatly affected by all levels of shade. In general, the results suggest that these hybrids are intermediate to the parental taxa in terms of shade tolerance.     

Structural and physiological responses of two invasive weeds, <Emphasis Type="Italic">Mikania micrantha</Emphasis> and <Emphasis Type="Italic">Chromolaena odorata</Emphasis>, to contrasting light and soil water conditions

To better understand the requirement of light and soil water conditions in the invasion sites of two invasive weeds, Mikania micrantha and Chromolaena odorata, we investigated their structural and physiological traits in response to nine combined treatments of light [full, medium and low irradiance (LI)] and soil water (full, medium and low field water content) conditions in three glasshouses. Under the same light conditions, most variables for both species did not vary significantly among different water treatments. Irrespective of water treatment, both species showed significant decreases in maximum light saturated photosynthetic rate (P _max), photosynthetic nitrogen-use efficiency, and relative growth rate under LI relative to full irradiance; specific leaf area, however, increased significantly from full to LI though leaf area decreased significantly, indicating that limited light availability under extreme shade was the critical factor restricting the growth of both species. Our results also indicated that M. micrantha performed best under a high light and full soil water combination, while C. odorata was more efficient in growth under a high light and medium soil water combination.     

Carbohydrate storage,survival, and growth of two evergreen <Emphasis Type="Italic">Nothofagus</Emphasis> species in two contrasting light environments

A number of traits have been attributed important roles in tolerance of shade by plants. Some explanations emphasize traits enhancing net carbon gain; others emphasize energy conservation traits such as storage of non-structural carbohydrates (NSC). To date, cross-species studies have provided mixed support for the role of NSC storage in low-light survival. We examined NSC status, survival, biomass, and growth of large seedlings of two evergreen species of differing shade tolerance (Nothofagus nitida and N. dombeyi) grown in deep shade and 50% light for two growing seasons. We expected to find higher NSC concentration in the more shade-tolerant N. nitida and since allocation to storage involves sacrificing growth, higher growth rate in the shade-intolerant N. dombeyi. NSC concentration of both species was >twofold higher in 50% light than in deep shade, and in roots and stems did not differ significantly between species in either environment. NSC contents per plant were also similar between dead and living plants in deep shade. N. dombeyi outgrew N. nitida in 50% light, while this pattern was reversed in deep shade. Survival in deep shade was not correlated with NSC concentration. Leaf mass fraction was similar between species in 50% light, but lower in N. dombeyi in deep shade. Results provide little evidence of a link between carbohydrate storage and low-light survival in Nothofagus species, and support the view that understorey survival is primarily a function of net carbon gain. Patterns of variation in NSC concentration of the temperate species we studied are likely dominated by more important influences than adaptation to shade, such as limitation of growth or adaptation to cold stress.