On the significance of belowground overyielding in temperate mixed forests: separating species identity and species diversity effects

Complementary soil exploration by the root systems of coexisting tree species has been hypothesised to result in a higher root biomass of mixed forests than of monocultures but the existing evidence for a belowground diversity effect in forests is scarce and not conclusive. In a species‐rich temperate broad‐leaved forest, we analysed the fine root biomass (roots ≤ 2 mm) and necromass in 100 plots differing in tree species diversity (one to three species) and species composition (all possible combinations of five species of the genera Acer, Carpinus, Fagus, Fraxinus and Tilia) which allowed us to separate possible species diversity and species identity effects on fine root biomass. We found no evidence of a positive diversity effect on standing fine root biomass and thus of overyielding in terms of root biomass. Root necromass decreased with increasing species diversity at marginal significance. Various lines of evidence indicate significant species identity effects on fine root biomass (10–20% higher fine root biomass in plots with presence of maple and beech than in plots with hornbeam; 100% higher fine root biomass in monospecific beech and ash plots than in hornbeam plots; differences significant). Ash fine roots tended to be over‐represented in the 2‐ and 3‐species mixed plots compared to monospecific ash plots pointing at apparent belowground competitive superiority of Fraxinus in this mixed forest. Our results indicate that belowground overyielding and spatial complementarity of root systems may be the exception rather than the rule in temperate mixed forests.     

Competition for nitrogen sources between European beech (Fagus sylvatica) and sycamore maple (Acer pseudoplatanus) seedlings

To investigate the short‐term consequences of direct competition between beech and sycamore maple on root N uptake and N composition, mycorrhizal seedlings of both tree species were incubated for 4 days (i.e. beech only, sycamore maple only or both together) in an artificial nutrient solution with low N availability. On the fourth day, N uptake experiments were conducted to study the effects of competition on inorganic and organic N uptake. For this purpose, multiple N sources were applied with a single label. Furthermore, fine roots were sampled and analysed for total amino acids, soluble protein, total nitrogen, nitrate and ammonium content. Our results clearly show that both tree species were able to use inorganic and organic N sources. Uptake of inorganic and organic N by beech roots was negatively affected in the presence of the competing tree species. In contrast, the presence of beech stimulated inorganic N uptake by sycamore maple roots. Both the negative effect of sycamore maple on N uptake of beech and the positive effect of beech on N uptake of sycamore maple led to an increase in root soluble protein in beech, despite an overall decrease in total N concentration. Thus, beech compensated for the negative effects of the tree competitor on N uptake by incorporating less N into structural N components, but otherwise exhibited the same strategy as the competitor, namely, enhancing soluble protein levels in roots when grown under competition. It is speculated that enhanced enzyme activities of so far unknown nature are required in beech as a defence response to inter‐specific competition.     

Effects of CO2 enrichment on whole-plant carbon budget of seedlings of Fagus grandifolia and Acer saccharum in low irradiance

Carbon exchange rates (CER) and whole-plant carbon balances of beech (Fagus grandifolia) and sugar maple (Acer saccharum) were compared for seedlings grown under low irradiance to determine the effects of atmospheric CO₂ enrichment on shade-tolerant seedlings of co-dominant species. Under contemporary atmospheric CO₂, photosynthetic rate per unit mass of beech was lower than for sugar maple, and atmospheric CO₂ enrich ment enhanced photosynthesis for beech only. Aboveground respiration per unit mass decreased with CO₂ enrichment for both species while root respiration per unitmass decreased for sugar maple only. Under contemporary atmoapheric CO₂, beech had lower C uptake per plant than sugar maple, while C losses per plant to nocturnal aboveground and root respiration were similar for both species. Under elevated CO₂, C uptake per plant was similar for both species, indicating a significant relative increase in whole-seedling CER with CO₂ enrich ment for beech but not for sugar maple. Total C loss per plant to aboveground respiration was decreased for beech only because increase in sugar maple leaf mass counterbalanced a reduction in respiration rates. Carbon loss to root respiration per plant was not changed by CO₂ enrichment for either species. However, changes in maintenance respiration cost and nitrogen level suggest changes in tissue composition with elevated CO₂. Beech had a greater net daily C gain with CO₂ enrichment than did sugar maple in contrast to a lower one under contemporary CO₂. Elevated CO₂ preferentially enhances the net C balance of beech by increasing photosynthesis and reducing respiration cost. In all cases, the greatest C lost was by roots, indicating the importance of belowground biomass in net C gain. Relative growth rate estimated from biomass accumulation was not affected by CO₂ enrichment for either species possibly because of slow growth under low light. This study indicates the importance of direct effects of CO₂ enrichment when predicting potential change in species distribution with global climate change.     

Rooting strategy of naturally regenerated beech in Silver birch and Scots pine woodlands

This study investigated the belowground development and strategy of late-successional European beech (Fagus sylvatica L.) in ageing natural Scots pine (Pinus sylvestris L.) and Silver birch (Betula pendula Roth.) woodlands in a French volcanic mid-elevation area. For this purpose root biomass, root profile and fine-root architecture of competitor trees were examined in 53 mixed pine–beech and 42 birch–beech woodlands along a stand maturation gradient, using the root auger technique (0–75-cm). The total beech fine-root biomass highly correlated with aerial dimensions such as stem height and girth, whereas it moderately correlated with its age, thus indicating the effects of competition. Basic stand biometric data such as stand density and basal area had no significant effect on beech root biomass. Conversely, competition indices taking into account the vertical dimensions of competitor trees were efficient, probably due to redundancy with beech height. At similar age and height, beeches under birch had a greater belowground development than beeches under pine. Each species exhibited specific rooting pattern and plasticity of fine-root architecture along the gradients of stand maturation and competition. Beech had a heart-shaped rooting habit in both mixings, which strongly increased along stand maturation. Its fine-root system adopted a foraging strategy to respond to increasing stand competition. The Scots pine fine-root system was plate-like and showed a low morphological plasticity, thus presumably a conservative strategy. Silver birch exhibited a high biomass and a foraging capacity in the topsoil but a loose root system in the subsoil. The coexistence of pine and beech roots in the upper soil presumably leads to a high belowground competition. Beech root system becomes predominant throughout the soil profile and it adopts an efficient foraging strategy, but at the expense of its belowground development. Conversely, the niche partitioning strategy between beech and birch may explain why beech develops strongly belowground in spite of the fact that birch has a dense rooting and a competitive fine-root architecture. As a consequence, beech mid-term regeneration and development may be facilitated under birch as compared with pine.     

Competition increasingly dominates the responsiveness of juvenile beech and spruce to elevated CO2 and/or O3 concentrations throughout two subsequent growing seasons

Saplings of Fagus sylvatica and Picea abies were grown in mono‐ and mixed cultures in a 2‐year phytotron study under all four combinations of ambient and elevated ozone (O₃) and carbon dioxide (CO₂) concentrations. The hypotheses tested were (1) that the competitiveness of beech rather than spruce is negatively affected by the exposure to enhanced O₃ concentrations, (2) spruce benefits from the increase of resource availability (elevated CO₂) in the mixed culture and (3) that the responsiveness of plants to CO₂ and O₃ depends on the type of competition (i.e. intra vs. interspecific). Beech displayed a competitive disadvantage when growing in mixture with spruce: after two growing seasons under interspecific competition, beech showed significant reductions in leaf gas exchange, biomass development and crown volume as compared with beech plants growing in monoculture. In competition with spruce, beech appeared to be nitrogen (N)‐limited, whereas spruce tended to benefit in terms of its plant N status. The responsiveness of the juvenile trees to the atmospheric treatments differed between species and was dominated by the type of competition: spruce growth benefited from elevated CO₂ concentrations, while beech growth suffered from the enhanced O₃ regime. In general, interspecific competition enhanced these atmospheric treatment effects, supporting our hypotheses. Significant differences in root : shoot biomass ratio between the type of competition under both elevated O₃ and CO₂ were not caused by readjustments of biomass partitioning, but were dependent on tree size. Our study stresses that competition is an important factor driving plant development, and suggests that the knowledge about responses of plants to elevated CO₂ and/or O₃, acquired from plants growing in monoculture, may not be transferred to plants grown under interspecific competition as typically found in the field.     

The relation between above- and belowground biomass allocation patterns and competitive ability

Summary In a 2-year experiment, the evergreen shrubsErica tetralix andCalluna vulgaris (dominant on nutrient-poor heathland soils) and the perennial deciduous grassMolinia caerulea (dominant on nutrient-rich heathland soils) were grown in replacement series in a factorial combination of four competition types (no competition, only aboveground competition, only belowground competition, full competition) and two levels of nutrient supply (no nutrients and 10 g N+2 g P+10 g K m⁻² yr⁻¹). Both in the unfertilized and in the fertilized treatmentsMolinia allocated about twice as much biomass to its root system than didErica andCalluna. In all three species the relative amount of biomass allocated to the roots was lower at high than at low nutrient supply. The relative decrease was larger forMolinia than forErica andCalluna. In the fertilized monocultures biomass of all three species exceeded that in the unfertilized series.Molinia showed the greatest biomass increase. In the unfertilized series no effects of interspecific competition on the biomass of each species were observed in either of the competition treatments. In the fertilized mixtures where only belowground competition was possibleMolinia increased its biomass at the expense of bothErica andCalluna. When only aboveground competition was possible no effects of interspecific competition on the biomass of the competing species were observed. However, in contrast with the evergreens,Molinia responded by positioning its leaf layers relatively higher in the canopy. The effects of full competition were similar to those of only belowground competition, so in the fertilized series belowground competition determined the outcome of competition. The high competitive ability ofMolinia at high nutrient supply can be attributed to the combination of (1) a high potential productivity, (2) a high percentage biomass allocation to the roots, (3) an extensive root system exploiting a large soil volume, and (4) plasticity in the spatial arrangement of leaf layers over its tall canopy. In the species under study the allocation patterns entailed no apparent trade-off between the abilities to compete for above- and belowground resources. This study suggests that this trade-off can be overcome by: (1) plasticity in the spatial arrangement of leaf layers and roots, and (2) compensatory phenotypic and species-specific differences in specific leaf area and specific root length.     

On the influence of provenance to soil quality enhanced stress reaction of young beech trees to summer drought

Climate projections propose that drought stress will become challenging for establishing trees. The magnitude of stress is dependent on tree species, provenance, and most likely also highly influenced by soil quality. European Beech (Fagus sylvatica) is of major ecological and economical importance in Central European forests. The species has an especially wide physiological and ecological amplitude enabling growth under various soil conditions within its distribution area in Central Europe. We studied the effects of extreme drought on beech saplings (second year) of four climatically distinct provenances growing on different soils (sandy loam and loamy sand) in a full factorial pot experiment. Foliar δ¹³C, δ¹⁵N, C, and N as well as above‐ and belowground growth parameters served as measures for stress level and plant growth. Low‐quality soil enhanced the effect of drought compared with qualitatively better soil for the above‐ and belowground growth parameters, but foliar δ¹³C values revealed that plant stress was still remarkable in loamy soil. For beeches of one provenance, negative sandy soil effects were clearly smaller than for the others, whereas for another provenance drought effects in sandy soil were sometimes fatal. Foliar δ¹⁵N was correlated with plant size during the experiment. Plasticity of beech provenances in their reaction to drought versus control conditions varied clearly. Although a general trend of declining growth under control or drought conditions in sandy soil was found compared to loamy soil, the magnitude of the effect of soil quality was highly provenance specific. Provenances seemed to show adaptations not only to drought but also to soil quality. Accordingly, scientists should integrate information about climatic pre‐adaptation and soil quality within the home range of populations for species distribution modeling and foresters should evaluate soil quality and climatic parameters when choosing donor populations for reforestation projects.     

Drought and recovery effects on belowground respiration dynamics and the partitioning of recent carbon in managed and abandoned grassland

The supply of soil respiration with recent photoassimilates is an important and fast pathway for respiratory loss of carbon (C). To date it is unknown how drought and land‐use change interactively influence the dynamics of recent C in soil‐respired CO₂. In an in situ common‐garden experiment, we exposed soil‐vegetation monoliths from a managed and a nearby abandoned mountain grassland to an experimental drought. Based on two ¹³CO₂ pulse‐labelling campaigns, we traced recently assimilated C in soil respiration during drought, rewetting and early recovery. Independent of grassland management, drought reduced the absolute allocation of recent C to soil respiration. Rewetting triggered a respiration pulse, which was strongly fuelled by C assimilated during drought. In comparison to the managed grassland, the abandoned grassland partitioned more recent C to belowground respiration than to root C storage under ample water supply. Interestingly, this pattern was reversed under drought. We suggest that these different response patterns reflect strategies of the managed and the abandoned grassland to enhance their respective resilience to drought, by fostering their resistance and recovery respectively. We conclude that while severe drought can override the effects of abandonment of grassland management on the respiratory dynamics of recent C, abandonment alters strategies of belowground assimilate investment, with consequences for soil‐CO₂ fluxes during drought and drought‐recovery.     

Root Dynamics of Cultivar and Non‐Cultivar Population Sources of Two Dominant Grasses during Initial Establishment of Tallgrass Prairie

Dominance of warm‐season grasses modulates tallgrass prairie ecosystem structure and function. Reintroduction of these grasses is a widespread practice to conserve soil and restore prairie ecosystems degraded from human land use changes. Seed sources for reintroduction of dominant prairie grass species include local (non‐cultivar) and selected (cultivar) populations. The primary objective of this study was to quantify whether intraspecific variation in developing root systems exists between population sources (non‐cultivar and cultivar) of two dominant grasses (Sorghastrum nutans and Schizachyrium scoparium) widely used in restoration. Non‐cultivar and cultivar grass seedlings of both species were isolated in an experimental prairie restoration at the Konza Prairie Biological Station. We measured above‐ and belowground net primary production (ANPP and BNPP, respectively), root architecture, and root tissue quality, as well as soil moisture and plant available inorganic nitrogen (N) in soil associated with each species and source at the end of the first growing season. Cultivars had greater root length, surface area, and volume than non‐cultivars. Available inorganic N and soil moisture were present in lower amounts in soil proximal to roots of cultivars than non‐cultivars. Additionally, soil NO₃–N was negatively correlated with root volume in S. nutans cultivars. While cultivars had greater BNPP than non‐cultivars, this was not reflected aboveground root structure, as ANPP was similar between cultivars and non‐cultivars. Intraspecific variation in belowground root structure and function exists between cultivar and non‐cultivar sources of the dominant prairie grasses during initial reestablishment of tallgrass prairie. Population source selection should be considered in setting restoration goals and objectives.     

Fine root morphological and functional traits in Fagus sylvatica and Fraxinus excelsior saplings as dependent on species, root order and competition

Aims

The mechanisms of belowground competition are not well understood. Addressing literature reports on competition-induced changes in tree fine root morphology, we conducted a growth experiment with tree saplings to investigate competition effects on important root morphological and functional traits in a root order-focused analysis.

Methods

European beech and European ash saplings were grown for 34 months in containers under greenhouse conditions in monoculture (2 conspecific plants), in mixture (1 beech and 1 ash) or as single plants. The root system was fractionated according to root orders and eight morphological and functional properties were determined.

Results

Root order was the most influential factor affecting the fine root traits (except for root diameter and δ¹³C); a significant species identity effect was found for root diameter, tissue density, N concentration and δ¹³C. Ash fine roots were thicker, but had lower tissue densities, contained more N and had systematically higher δ¹³C values than beech roots. The competition treatments had no significant effect on morphological root traits but altered δ¹³C in the 2nd root order.

Conclusion

Neither intra- nor interspecific root competition affected fine root morphology significantly suggesting that competition-induced root modification may not be a universal phenomenon in temperate trees.     

Interactions between root and shoot competition vary among species

Understanding how the competition varies with productivity is essential for differentiating among alternative models of plant community organization. Prior attempts to explain shifts in root and shoot competition along gradients have generally assumed an additive interaction between the two competitive forms, using an experimental design which does not fully separate both above‐ and belowground processes. At the most basic level, few field studies have separated root and shoot competition, and we have limited knowledge about both the relative importance of these processes, and how they interact to affect plant growth in the field. Presented here are findings from a field study in which root and shoot competition were experimentally separated by using root exclusion tubes and neighbor tiebacks in an early successional community. Individuals of four species (Abutilon theophrasti, Amaranthus retroflexus, Rumex crispus, and Plantago lanceolata) were grown at two levels of fertilization with full competition, aboveground competition only, belowground competition only, or neither above‐ nor belowground competition. Competition was measured as competitive response, which is the natural log of the relative biomass of a target plant grown with competition compared to growth without competition. In contrast to predictions from current models of productivity‐competition relationships, but in agreement with other experimental studies, there was no change in the strengths or root, shoot, or total competition with a modest increase in productivity. Despite no effect of fertilization on the strength of competition, the form of interaction between root and shoot competition varied both as a function of species identity and fertilization. For both of the rosette forming species, the combined effects of root and shoot competition were less than predicted assuming no interaction (a “negative interaction”), with one species switching from a negative to an additive interaction with fertilization. The fact that fertilization caused a shift in the root‐shoot interaction, but not in the total strength of root and shoot competition, suggests that the root‐shoot interaction is itself a highly labile variable. If root‐shoot interactions are common in natural systems, then simply measuring the strength of one form of competition in no way provides any information about the overall importance of that competitive form to plant growth.     

Effect of soil nutrients,neighbor identities and root separation types on intra‐ and interspecific interaction among three clonal plant species

In natural environments, plants frequently interact with both heterospecific and conspecific neighbors. The intensity of belowground plant interaction with neighboring species commonly varies with the availability of soil nutrients in the habitats. According to classical ecological theory, competition between conspecific neighbors may be more severe than competition between unrelated species due to the similar nutrient requirements of close relatives, especially when nutrients are scarce in the habitat. However, many recent studies have shown the opposite pattern, and suggested an alternative mechanism based on species recognition. Taking Zoysia sinica as the focal species, we conducted a controlled experiment to test the results of intraspecific and interspecific interactions among three clonal species Zoysia sinica, Zoysia japonica and Alternanthera philoxeroides, which represent a conspecific, a close relative and a distant relative of the focal species, respectively, and at different root treatments (no separation NS, clone separation CS and ramet separation RS) and two nutrient levels. The results showed that Z. sinica recognized conspecific plants in the NS and CS treatments, and did not show above or belowground competition with these. The performance of the focal plant (Z. sinica) was better when it was grown with a conspecific neighbor as compared to all other types of neighbors. In all root separation treatments, the competition was more intense when Z. sinica grew with a close relative (Z. japonica) than when growing with a distant relative (A. philoxeroides). Generally, competition between plants was more intense at the high nutrient level than at the low nutrient level, suggesting that both soil nutrients and a species recognition mechanism play a significant role for the intra‐ and interspecific interaction and fitness of these three neighboring clonal species.     

Influence of water availability on competitive interactions between plant species on sandy soils

The gravimetric soil water content of four different successional stages of inland dune vegetation revealed pronounced seasonal changes and particularly a major drought period in the late summer. In order to analyze the effect of these seasonal water changes on the competitive interactions of two dominant species (Carex arenaria andDeschampsia flexuosa) in one of these stages root development was compared under experimentally manipulated water conditions in a minirhizotron experiment. Under well-watered conditions, the root growth ofC. arenaria andD. flexuosa was not influenced by competition, but was significantly reduced forC. arenaria at low water availability. Accordingly, the relative yield based on total biomass was significantly lower under water shortage than under well-watered conditions forC. arenaria, indicating higher interspecific competition compared to intraspecific competitive interactions. These results indicate (i) that soil water status is an important factor for belowground competitive interactions and (ii) that seasonal changes in soil water content may have a pronounced effect on the outcome of competition and, in consequence, on the structure of the respective plant communities.     

Competitive interactions shape plant responses to nitrogen fertilization and drought: evidence from a microcosm experiment with Lilium bulbiferum L. and Secale cereale L.

Many recent studies have analysed plant species responses to environmental change, but interactive effects of global change drivers and how they are modulated by biotic interactions are still poorly understood. In a mesocosm experiment, we studied the interactive effects of nitrogen (N) fertilization and drought events on plant growth and how these effects are shaped by competitive interactions, using a segetal plant community typical of the lowlands of central Europe (composed of Lilium bulbiferum (segetal species) and Secale cereale (crop species)). We expected that N fertilization increases the drought sensitivity of Lilium (negative interaction effect), and that these effects are shaped by interspecific competition with Secale. Secale and Lilium showed opposing responses to N fertilization (second year of the experiment): Whilst Secale aboveground and belowground biomass almost doubled with N fertilization, Lilium aboveground and belowground biomass showed no response or decreased, respectively, providing Secale with a competitive advantage. Lilium aboveground tissue dieback (as a proxy for growth vigour) was 22% in N and 35% in drought treatments (control: 6%), but reached 91% when combining these treatments. Increasing Lilium tissue dieback was strongly related to decreasing belowground (root) biomass, caused by both negative direct effects of combined treatments (N fertilization?+?drought), and negative indirect effects acting via treatment-induced increase in Secale biomass. Our results demonstrate that competitive interactions can shape the effects of global change drivers on plant growth. This knowledge in turn could be important for plant species conservation, particularly in the face of ongoing shifts in environmental conditions.

     

To compete or not to compete: an experimental study of interactions between plant species with contrasting root behaviour

Game-theoretic models predict that plants with root systems that avoid belowground competition will be displaced by plants that overproduce roots in substrate shared with competitors. Despite this, both types of root response to neighbours have been documented. We used two co-occurring clonal species (Glechoma hederacea and Fragaria vesca) with contrasting root responses to neighbours (avoidance of competition and contesting of resources, respectively) to examine whether functional variation in other traits affected the success of each rooting strategy, leading to a different outcome from that predicted on the basis of root behaviour alone. Vegetative propagation rates, morphology and biomass allocation patterns were examined when each species was challenged with competition from physically separate ramets with either the same rooting strategy (intraclonal competition) or the contrasting rooting strategy (interspecific competition). Contrary to the predictions of game-theoretic models, the species that exhibits avoidance of root competition (Glechoma) was not competitively inferior to the species that does not (Fragaria). Glechoma achieved greater total mass in the interspecific treatment than in the intraclonal treatment. However, Fragaria did not experience more intense competition from Glechoma than it did in the intraclonal treatment. Strong interference between the two species appeared to be avoided because Glechoma invested preferentially in rapid exploitation of unoccupied space, whereas Fragaria invested in increasing the competitive ability and local persistence of established ramets. Our results suggest that interspecific trade-offs between traits related to competitive ability and resource exploitation can allow coexistence of species with contrasting rooting behaviours. Full assessment of the adaptive value of different root responses to neighbours therefore requires concurrent consideration of the combined effects of a wide array of functional traits.     

The symmetry of competitive interactions in mixed Norway spruce,silver fir and European beech forests

Questions

We aim for a better understanding of the different modes of intra‐ and inter‐specific competition in two‐ and three‐species mixed‐forests. How can the effect of different modes of competitive interactions be detected and integrated into individual tree growth models? Are species interactions in spruce–fir–beech forests more associated with size‐symmetric or size‐asymmetric competition? Do competitive interactions between two of these species change from two‐ to three‐species mixtures?

Location

Temperate mixed‐species forests in Central Europe (Switzerland).

Methods

We used data from the Swiss National Forest Inventory to fit basal area increment models at the individual tree level, including the effect of ecological site conditions and indices of size‐symmetric and size‐asymmetric competition. Interaction terms between species‐specific competition indices were used to disentangle significant differences in species interactions from two‐ to three‐species mixtures.

Results

The growth of spruce and fir was positively affected by increasing proportions of the other species in spruce–fir mixtures, but negative effects were detected with increasing presence of beech. We found that competitive interactions for spruce and fir were more related to size‐symmetric competition, indicating that species interactions might be more associated with competition for below‐ground resources. Under constant amounts of stand basal area, the growth of beech clearly benefited from the increasing admixture of spruce and fir. For this species, patterns of size‐symmetric and size‐asymmetric competitive interactions were similar, indicating that beech is a strong self‐competitor for both above‐ground and below‐ground resources. Only for silver fir and beech, we found significant changes in species interactions from two‐ to three‐species mixtures, but these were not as prominent as the effects due to differences between intra‐ and inter‐specific competition.

Conclusions

Species interactions in spruce–fir–beech, or other mixed forests, can be characterized depending on the mode of competition, allowing interpretations of whether they occur mainly above or below ground level. Our outcomes illustrate that species‐specific competition indices can be integrated in individual tree growth functions to express the different modes of competition between species, and highlight the importance of considering the symmetry of competition alongside competitive interactions in models aimed at depicting growth in mixed‐species forests.

     

Effects of land‐use changes on ecosystem services: decrease in ant predation in human‐dominated landscapes in central Brazil

Anthropogenic disturbances often affect the abundance and diversity of ants (Hymenoptera: Formicidae) but relatively few studies have explored the implications of such changes on the ecosystem services mediated by these insects. Here, we evaluated how the transformation of Cerrado savanna habitats into crop plantations affects the abundance, diversity, and the predatory activity of ants. A survey of the ant faunas foraging above‐ and belowground was performed in six crop and six non‐crop (i.e., native vegetation) sites. Above‐ and belowground rates of ant predation were estimated at these same sites using mealworms, Tenebrio molitor L. (Coleoptera: Tenebrionidae), as baits, simulating crop herbivores. Belowground predation rates were significantly greater in the non‐crop sites, despite the lack of difference in overall abundance and species richness of ants foraging belowground between the crop vs. non‐crop sites. In contrast, we did not detect any significant difference in aboveground predation rates between crop vs. non‐crop sites even though there were significantly more species of ants foraging aboveground in the non‐crop sites. Army ants (subfamily Dorylinae) were the main predatory species belowground, and their abundance was significantly greater in non‐crop sites. In contrast, the main predators aboveground were omnivore ants of the genera Pheidole and Solenopsis, which had similar abundances in the crop and non‐crop sites. Overall, our results indicate that transformation of native Cerrado habitats into crop plantations reduces the abundance of some important predatory species, notably those that forage belowground, and this may negatively affect the potential for ants to provide pest control services in agroecosystems.     

A California grasslands alkali specialist,Hemizonia pungens ssp. pungens,prefers non‐alkali soil

Question: Edaphically severe habitats commonly support edaphic endemics, specialized plant species that do not occur elsewhere. The endemism of native plant species in edaphically specialized habitats suggests either (a) that these native endemic species are uniquely specialized to survive and grow better under the conditions prevalent in these harsh areas, or (b) that these areas represent refuges from competition with other (often exotic) species. Location: Central Valley, California, USA. Methods: We surveyed the vegetation distribution in alkali sinks and carried out a reciprocal transplant greenhouse experiment crossed with a competition treatment to examine the interplay between interspecific competition and edaphic factors in determining relative performance of an alkali endemic forb (Hemizonia pungens ssp. pungens) and its exotic grass competitor (Lolium multiflorum). Results: Lolium consistently performed better in non‐alkali soil. In contrast, Hemizonia, in the absence of competition, performed significantly better on the non‐alkali soils, but in competition with Lolium, performed equally well on alkali and non‐alkali soils. Conclusions: These results suggest that Hemizonia does not inherently prefer harsh alkali soil, but is better able to tolerate alkali soil and may be excluded from more moderate soils by competition. Therefore, edaphic and biotic effects may interact to determine the spatial distribution of this edaphic endemic. Our data suggest a mechanism by which competition between native and exotic species on impoverished soils leads to dominance of native species and creation of refugia for native species where exotic species are unable to thrive.     

Effects of increasing root carbon investment on the mortality and resprouting of Haloxylon ammodendron seedlings under drought

• Tree mortality induced by drought is one of the most complex processes in ecology. Although two mechanisms associated with water and carbon balance are proposed to explain tree mortality, outstanding problems still exist.
• Here, in order to test how the root system benefits survival and resprouting of Haloxylon ammodendron seedlings, we examined the various water‐ and carbon‐related physiological indicators (shoot water potential, photosynthesis, dark respiration, hydraulic conductance and non‐structural carbohydrates [NSC]) of H. ammodendron seedlings, which were grown in drought and control conditions throughout a grow season in greenhouse.
• The survival time of the seedling root system (died 70 days after drought) doubled the survival time of the shoot (died at 35 days). Difference in survival time between shoot and root resulted from sustained root respiration supported by increased NSC in roots under drought. Furthermore, investment into the root contributed to resprouting following drought. Based on these results, a death criterion is proposed for this species. The time sequence of major events indicated that drought shifted carbon allocation between shoot and root and altered the flux among different sinks (growth, respiration or storage). The interaction of water and carbon processes determined death or survival of droughted H. ammodendron seedlings.
• These findings revealed that the ‘root protection’ strategy is critical in determining survival and resprouting of this species, and provided insights into the effects of carbon and water dynamics on tree mortality.

     

Interspecific competition enhances nitrogen fixation in an actinorhizal shrub

In forest understory restoration, the establishment of reintroduced species may be strongly linked to their ability to compete for belowground resources. In this study, we provide isotopic and morphological evidence for competition-induced increases in nitrogen fixation by Morella cerifera (L.) Small (wax myrtle) when planted with Pinus palustris Mill (longleaf pine). Compared to a competition-free treatment, we found no significant differences in tissue N concentrations for M. cerifera. However, ¹⁵N enrichment in leaves, stems and roots, as well as whole-plant values for nitrogen derived from fertilizer were significantly lower when the plants were subject to interspecific competition from P. palustris. Plants in the competition treatment also allocated a significantly greater percentage of belowground biomass to root nodules than those in the competition-free treatment (0.65 vs. 0.41%). This strongly suggests that M. cerifera is capable of upregulating nitrogen fixation in response to interspecific competition. This may help explain why M. cerifera outperformed non-nitrogen-fixing species reintroduced on the same site.