Meta-analysis reveals that the effects of precipitation change on soil and litter fauna in forests depend on body size.

Anthropogenic climate change is altering precipitation regimes at a global scale. While precipitation changes have been linked to changes in the abundance and diversity of soil and litter invertebrate fauna in forests, general trends have remained elusive due to mixed results from primary studies. We used a meta-analysis based on 430 comparisons from 38 primary studies to address associated knowledge gaps, (i) quantifying impacts of precipitation change on forest soil and litter fauna abundance and diversity, (ii) exploring reasons for variation in impacts and (iii) examining biases affecting the realism and accuracy of experimental studies. Precipitation reductions led to a decrease of 39% in soil and litter fauna abundance, with a 35% increase in abundance under precipitation increases, while diversity impacts were smaller. A statistical model containing an interaction between body size and the magnitude of precipitation change showed that mesofauna (e.g. mites, collembola) responded most to changes in precipitation. Changes in taxonomic richness were related solely to the magnitude of precipitation change. Our results suggest that body size is related to the ability of a taxon to survive under drought conditions, or to benefit from high precipitation. We also found that most experiments manipulated precipitation in a way that aligns better with predicted extreme climatic events than with predicted average annual changes in precipitation and that the experimental plots used in experiments were likely too small to accurately capture changes for mobile taxa. The relationship between body size and response to precipitation found here has far-reaching implications for our ability to predict future responses of soil biodiversity to climate change and will help to produce more realistic mechanistic soil models which aim to simulate the responses of soils to global change.

Effects of litter quality on foraging behaviour and demographic parameters in Folsomia candida (Collembola).

Litter quality has long been associated with demographic parameters of Collembola populations. However, little is known about the capacity of Collembola to perceive and seek better litter quality. To address this gap, three complementary laboratory experiments were carried out with the Collembola Folsomia candida. First, populations were fed on three different types of leaf litters (Quercus pubescens, Acer opalus and Prunus avium) and a control (agar-agar-brewer's yeast mixture) for 6 weeks to assess their impacts on demography (reproduction rate and population size). Second, the body length of individuals differentially fed with the same four types of resources was measured to assess a functional trait that can potentially affect movement parameters such as prospected area or foraging speed. Third, F. candida single individuals were exposed to the same litter quality gradient and placed at an increasing distance from the litter (from 1 to 5 cm). For 10 min, their foraging behaviour was recorded which included prospected area, foraging speed, perception distance and success in reaching the litter (foraging success). As expected, low-quality litter (i.e. Q. pubescens) contributed to low population growth compared to the control treatment and the high-quality litters (P. avium and A. opalus). In the third experiment, the probability of finding the resource was negatively correlated to the distance, but was unrelated to the litter quality and the Collembola body length. When resource was perceived, F. candida was able to switch from non-directional to directional movements, with a large variability in the perception distance from a few millimetres to several centimetres. Taken together, our results indicate that litter quality plays a relevant role in Collembola demographic parameters once the population settles on litter patch, but not on foraging behaviour to select high-quality resources.

Amplified Drought Alters Leaf Litter Metabolome, Slows Down Litter Decomposition, and Modifies Home Field (Dis)Advantage in Three Mediterranean Forests.

In Mediterranean ecosystems, the projected rainfall reduction of up to 30% may alter plant-soil interactions, particularly litter decomposition and Home Field Advantage (HFA). We set up a litter transplant experiment in the three main forests encountered in the northern part of the Medi-terranean Basin (dominated by either Quercus ilex, Quercus pubescens, or Pinus halepensis) equipped with a rain exclusion device, allowing an increase in drought either throughout the year or concentrated in spring and summer. Senescent leaves and needles were collected under two precipitation treatments (natural and amplified drought plots) at their "home" forest and were left to decompose in the forest of origin and in other forests under both drought conditions. MS-based metabolomic analysis of litter extracts combined with multivariate data analysis enabled us to detect modifications in the composition of litter specialized metabolites, following amplified drought treatment. Amplified drought altered litter quality and metabolomes, directly slowed down litter decomposition, and induced a loss of home field (dis)advantage. No indirect effect mediated by a change in litter quality on decomposition was observed. These results may suggest major alterations of plant-soil interactions in Mediterranean forests under amplified drought conditions.

Amplified drought induced by climate change reduces seedling emergence and increases seedling mortality for two Mediterranean perennial herbs.

Seedling recruitment is a bottleneck for population dynamics and range shift. The vital rates linked to recruitment by seed are impacted by amplified drought induced by climate change. In the Mediterranean region, autumn and winter seedling emergence and mortality may have strong impact on the overall seedling recruitment. However, studies focusing on the temporal dynamic of recruitment during these seasons are rare. This study was performed in a deciduous Mediterranean oak forest located in southern France and quantifies the impact of amplified drought conditions on autumn and winter seedling emergence and seedling mortality rates of two herbaceous plant species with meso-Mediterranean and supra-Mediterranean distribution (respectively, Silene italica and Silene nutans). Seedlings were followed from October 2019 to May 2020 in both undisturbed and disturbed plots where the litter and the aboveground biomass have been removed to create open microsites. Amplified drought conditions reduced seedling emergence and increased seedling mortality for both Silene species but these negative effects were dependent on soil disturbance conditions. Emergence of S. italica decreased only in undisturbed plots (-7%) whereas emergence of S. nutans decreased only in disturbed plots (-10%) under amplified drought conditions. The seedling mortality rate of S. italica was 51% higher under amplified drought conditions in undisturbed plots while that of S. nutans was 38% higher in disturbed plots. Aridification due to lower precipitation in the Mediterranean region will negatively impact the seedling recruitment of these two Silene species. Climate change effects on early vital rates may likely have major negative impacts on the overall population dynamic.

Volatilome of Aleppo Pine litter over decomposition process.

Biogenic Volatile Organic Compounds (BVOC) are largely accepted to contribute to both atmospheric chemistry and ecosystem functioning. While the forest canopy is recognized as a major source of BVOC, emissions from plant litter have scarcely been explored with just a couple of studies being focused on emission patterns over litter decomposition process. The aim of this study was to quantitatively and qualitatively characterize BVOC emissions (C1-C15) from Pinus halepensis litter, one of the major Mediterranean conifer species, over a 15-month litter decomposition experiment. Senescent needles of P. halepensis were collected and placed in 42 litterbags where they underwent in situ decomposition. Litterbags were collected every 3 months and litter BVOC emissions were studied in vitro using both online (PTR-ToF-MS) and offline analyses (GC-MS). Results showed a large diversity of BVOC (58 compounds detected), with a strong variation over time. Maximum total BVOC emissions were observed after 3 months of decomposition with 9.18 µg gDM -1 hr-1 mainly composed by terpene emissions (e.g., α-pinene, terpinolene, ß-caryophyllene). At this stage, methanol, acetone, and acetic acid were the most important nonterpenic volatiles representing, respectively, up to 26%, 10%, and 26% of total emissions. This study gives an overview of the evolution of BVOC emissions from litter along with decomposition process and will thus contribute to better understand the dynamics and sources of BVOC emission in Mediterranean pine forests.

Legacy effect of green manure crops fertilized with calcium phosphite on maize production and soil properties.

Recycling phosphorus (P) is crucial to meet future P demand for crop production. We investigated the possibility to use calcium phosphite (Ca-Phi) waste, an industrial by-product, as P fertilizer following the oxidation of phosphite (Phi) to phosphate (Pi) during green manure (GM) cropping in order to target P nutrition of subsequent maize crop. In a greenhouse experiment, four GM crops were fertilized (38 kg P ha-1) with Ca-Phi, triple super phosphate (TSP) or without P (Control) in sandy and clay soils. The harvested GM biomass (containing Phi after Ca-Phi fertilization) was incorporated into the soil before maize sowing. Incorporation of GM residues containing Phi slowed down organic carbon mineralization in clay soil and mass loss of GM residues in sandy soil. Microbial enzymatic activities were affected by Ca-Phi and TSP fertilization at the end of maize crop whereas microbial biomass was similarly influenced by TSP and Ca-Phi in both soils. Compared to Control, Ca-Phi and TSP increased similarly the available P (up to 5 mg P kg-1) in sandy soil, whereas in clay soil available P increased only with Ca-Phi (up to 6 mg P kg-1), indicating that Phi oxidation occurred during GM crops. Accordingly, no Phi was found in maize biomass. However, P fertilization did not enhance aboveground maize productivity and P export, likely because soil available P was not limiting. Overall, our results indicate that Ca-Phi might be used as P source for a subsequent crop since Phi undergoes oxidation during the preliminary GM growth.

Valorization of calcium phosphite waste as phosphorus fertilizer: Effects on green manure productivity and soil properties.

The potential to use calcium phosphite (Ca-Phi) as phosphorus (P) fertilizer may represent an effective recycling of P-containing by-products. A greenhouse experiment was conducted to investigate the effect of Ca-Phi (38 kg P ha-1) on soil properties and the growth parameters of four green manure species in clay and sandy soils using Ca-Phi, TSP (triple superphosphate) and control (no fertilization) as treatments. Eight weeks after sowing, we measured aboveground biomass yield, phosphite (Phi) concentration in plant biomass, different soil P pools as well as microbial biomass nutrients. Compared to control, the addition of Ca-Phi did not negatively affect green manure yield, except for lupine (Lupinus albus L.) in clay soil. The Phi concentration in plant biomass varied across species and soil type with a maximum concentration of about 400 mg Phi kg-1 for mustard (Brassica juncea L.) in clay soil. Compared to control, TSP and Ca-Phi fertilization had a similar effect on different P pools and microbial biomass nutrients (C, N and P) although the response was soil-type dependent. In the sandy soil, after Ca-Phi addition the amount of available P (PNHCO3) increased to the same extent as in the TSP treatment (i.e. around 6 mg P kg-1) suggesting that Ca-Phi was, at least partly, oxidized. In the clay soil with high P fixing capacity, Ca-Phi promoted higher PNaHCO3 than TSP likely due to different solubility of chemical P forms. Additional studies are however required to better understand soil microbial responses and to quantify the P agronomical efficiency for the following crop under Ca-Phi fertilization.

Shift in plant-soil interactions along a lakeshore hydrological gradient.

Wetlands occupy the transitional zone between aquatic and terrestrial systems. Hydrological conditions have significant influence on wetland plant communities and soil biogeochemistry. However, our knowledge about plant-soil interactions in wetlands along hydrological gradients is still limited, although it is crucial to guide wetland management decisions and to adapt, whenever possible, hydrological conditions to the different plant communities. To this aim, we related vegetation composition, plant functional traits, soil physicochemical properties, soil microbial biomass, and soil enzymatic activities in wetlands on the southeastern shore of Neuchâtel lake, Switzerland, a lake whose level is partly regulated. Aboveground and belowground plant biomass and correspondent C, N and P concentrations remained constant or decreased moving from the vegetation community subjected to more frequent flooding events to the community with almost no flooding. The soil organic layer exhibited always higher nutrient concentrations and greater enzymatic activities than the organo-mineral and mineral layers. The chemical and biological characteristics of the soil organic layer showed decreasing values for most of the parameters along the hydrological gradient from lakeshore to upland wetland communities. On the basis of nutrient stoichiometry, plant-soil system in the plant community with most flooding events had no-nutrient limitation, while there was a N limitation in the transitional community. In the upland plant community where there was no flooding effect, the plant-soil system was characterized by N and P co-limitation. These findings are important because they provide a threshold for flooding regime by the lake in the context of optimization of lake level regulation under various stakeholders needs.

Seasonal variability of groundwater level effects on the growth of Carex cinerascens in lake wetlands.

Groundwater level is crucial for wetland plant growth and reproduction, but the extent of its effect on plant growth can vary along with changed precipitation and temperature at different seasons. In this context, we investigated the effect of two groundwater levels (10 cm vs. 20 cm depth) on growth and reproductive parameters of Carex cinerascens, a dominant plant species in the Poyang Lake wetland, during three seasons (spring, summer, and autumn) and during two consecutive years (2015 and 2016). Carex cinerascens showed low stem number, height, and individual and population biomass in summer compared to spring and autumn. 10 cm groundwater level was overall more suitable for plant growth resulting in higher stem height and biomass. However, the interactive effect between groundwater level and season clearly demonstrated that the effect of groundwater level on plant growth occurred mainly in autumn. After the withering of the plant population in summer, we observed that C. cinerascens growth recovered in autumn to similar values observed in spring only with 10 cm groundwater level. Consequently, we could deduce that lowering groundwater level in the studied Poyang Lake wetland will negatively impact C. cinerascens regeneration and growth particularly during the second growth cycle occurring in autumn. Additionally, our results showed that, independently of the season and groundwater level, population biomass of C. cinerascens was lower during drier year. Altogether, our findings suggest that water limitation due to both reduction in precipitation and decreased groundwater level during the year can strongly impact plant communities.

Allelopathic effects of volatile organic compounds released from Pinus halepensis needles and roots.

The Mediterranean region is recognized as a global biodiversity hotspot. However, over the last decades, the cessation of traditional farming in the north part of the Mediterranean basin has given way to strong afforestation leading to occurrence of abandoned agricultural lands colonized by pioneer expansionist species like Pinus halepensis. This pine species is known to synthesize a wide range of secondary metabolites, and previous studies have demonstrated strong allelopathic potentialities of its needle and root leachates. Pinus halepensis is also recognized to release significant amounts of volatile organic compounds (VOC) with potential allelopathic effects that have never been investigated. In this context, the objectives of the present study were to improve our knowledge about the VOC released from P. halepensis needles and roots, determine if these VOC affect the seed germination and root growth of two herbaceous target species (Lactuca sativa and Linum strictum), and evaluate if soil microorganisms modulate the potential allelopathic effects of these VOC. Thirty terpenes were detected from both, needle and root emissions with ß-caryophyllene as the major volatile. Numerous terpenes, such as ß-caryophyllene, δ-terpinene, or α-pinene, showed higher headspace concentrations according to the gradient green needles < senescent needles < needle litter. Seed germination and root growth of the two target species were mainly reduced in presence of P. halepensis VOC. In strong contrast with the trend reported with needle leachates in literature, we observed an increasing inhibitory effect of P. halepensis VOC with the progress of needle physiological stages (i.e., green needle < senescent needle < needle litter). Surprisingly, several inhibitory effects observed on filter paper were also found or even amplified when natural soil was used as a substrate, highlighting that soil microorganisms do not necessarily limit the negative effects of VOC released by P. halepensis on herbaceous target species.

Sponge inventory of the French Mediterranean waters, with an emphasis on cave-dwelling species.

Mediterranean sponges represent about 10 % of the world sponge biodiversity, with these sessile organisms dominating in terms of diversity and biomass in most of the rocky bottoms shaded from light. After 60 years of intensive study of the sponge diversity along the French coast, we present the first comprehensive reference-list for this biogeographic area. A total of 389 sponge species was recorded, of which 222 known in the Marseille region. In this area, special attention was paid to species from underwater caves. Although this particular habitat appeared as one of the richest, a wealth of hidden diversity still requires description. About 37 % of the sponge diversity can be found in underwater caves, most of these species being also distributed in other habitats. However, 23 % of this sponge diversity is cave-exclusive. An easy and rapid assessment method was developed with a selection of 65 representative sponge species, for the monitoring of semi-dark cave communities. This method, based on data acquisition with photoquadrats and their processing using a DataBase built with ACCESS, was deployed in 13 studied sites. Altogether, this study represents a useful contribution for marine environment managers who might refer to this French reference list and apply the rapid and easy assessment method in the framework of several European Directives and international Conventions.

Macroinvertebrate identity mediates the effects of litter quality and microbial conditioning on leaf litter recycling in temperate streams.

Plant litter decomposition is an essential ecosystem function that contributes to carbon and nutrient cycling in streams. Aquatic shredders, mainly macroinvertebrates, can affect this process in various ways; they consume leaf litter, breaking it down into fragments and creating suitable habitats or resources for other organisms through the production of fine particulate organic matter (FPOM). However, measures of litter-feeding traits across a wide range of aquatic macroinvertebrates are still rare. Here, we assessed the contributions of 11 species of freshwater macroinvertebrates to litter decomposition, by measuring consumption rate, FPOM production, and assimilation rate of highly decomposable (Alnus glutinosa) or poorly decomposable (Quercus robur) leaf litter types. In general, an increase in the quality of litter improved the litter consumption rate, and fungal conditioning of the leaf litter increased both the litter consumption rate and FPOM production. Macroinvertebrates specializing in leaf litter consumption also appeared to be the most sensitive to shifts in litter quality and the conditioning process. Contrary to expectations, the conditioning process did not increase the assimilation of low-quality litter. There was a strong correlation between the relative consumption rate (RCR) of the two litter types, and the relative FPOM production (RFP) was strongly correlated to the RCR. These findings suggest a consistent relationship between RCR and macroinvertebrate identity that is not affected by litter quality, and that the RFP could be inferred from the RCR. The varying responses of the macroinvertebrate feeding traits to litter quality and the conditioning process suggest that the replacement of a shredder invertebrate species by another species could have major consequences for the decomposition process and the detritus-based food web in streams. Further studies onto the importance of invertebrate identity and the effects of litter quality in a variety of freshwater ecosystems are needed to understand the whole ecosystem functioning and to predict its response to environmental changes.

Predator-prey interactions in a changing world: humic stress disrupts predator threat evasion in copepods.

Increasing inputs of colored dissolved organic matter (cDOM), which is mainly composed of humic substances (HS), are a widespread phenomenon of environmental change in aquatic ecosystems. This process of brownification alters the chemical conditions of the environment, but knowledge is lacking of whether elevated cDOM and HS levels interfere with the ability of prey species to evade chemical predator cues and thus affect predator-prey interactions. We assessed the effects of acute and prolonged exposure to HS at increasing concentrations on the ability of freshwater zooplankton to avoid predator threat (imposed by fish kairomones) in laboratory trials with two calanoid copepods (Eudiaptomus gracilis and Heterocope appendiculata). Populations of both species clearly avoided water containing fish kairomones. However, the avoidance behavior weakened with increasing HS concentration, suggesting that HS affected the ability of copepods to perceive or respond to the predator cue. The behavioral responses of the two copepod populations to increasing HS concentrations differed, with H. appendiculata being more sensitive than E. gracilis in an acute exposure scenario, whereas E. gracilis responded more strongly after prolonged exposure. Both showed similar physiological impairment after prolonged exposure, as revealed by their oxidative balance as a stress indicator, but mortality increased more strongly for H. appendiculata when the HS concentration increased. These results indicate that reduced predator threat evasion in the presence of cDOM could make copepods more susceptible to predation in future, with variation in the strength of responses among populations leading to changes in zooplankton communities and lake food-web structure.

The Impact of Competition and Allelopathy on the Trade-Off between Plant Defense and Growth in Two Contrasting Tree Species.

In contrast to plant-animal interactions, the conceptual framework regarding the impact of secondary metabolites in mediating plant-plant interference is currently less well defined. Here, we address hypotheses about the role of chemically-mediated plant-plant interference (i.e., allelopathy) as a driver of Mediterranean forest dynamics. Growth and defense abilities of a pioneer (Pinus halepensis) and a late-successional (Quercus pubescens) Mediterranean forest species were evaluated under three different plant interference conditions: (i) allelopathy simulated by application of aqueous needle extracts of Pinus, (ii) resource competition created by the physical presence of a neighboring species (Pinus or Quercus), and (iii) a combination of both allelopathy and competition. After 24 months of experimentation in simulated field conditions, Quercus was more affected by plant interference treatments than was Pinus, and a hierarchical response to biotic interference (allelopathy < competition < allelopathy + competition) was observed in terms of relative impact on growth and plant defense. Both species modulated their respective metabolic profiles according to plant interference treatment and thus their inherent chemical defense status, resulting in a physiological trade-off between plant growth and production of defense metabolites. For Quercus, an increase in secondary metabolite production and a decrease in plant growth were observed in all treatments. In contrast, this trade-off in Pinus was only observed in competition and allelopathy + competition treatments. Although Pinus and Quercus expressed differential responses when subjected to a single interference condition, either allelopathy or competition, species responses were similar or positively correlated when strong interference conditions (allelopathy + competition) were imposed.

Allelochemicals of Pinus halepensis as drivers of biodiversity in Mediterranean open mosaic habitats during the colonization stage of secondary succession.

The Mediterranean region is recognized as a global biodiversity hotspot. However, over the last 50 years or so, the cessation of traditional farming has given way to strong afforestation at the expense of open habitats. Pinus halepensis Miller, known to synthesize a wide range of secondary metabolites, is a pioneer expansionist species colonizing abandoned agricultural land that present high species richness. Here, laboratory bioassays were used to study the potential impact of P. halepensis on plant diversity through allelopathy, and the role of microorganisms in these interactions. Germination and growth of 12 target species naturally present in fallow farmlands were tested according to concentration of aqueous extracts obtained from shoots of young pines (aged about 5 years), with or without the presence of soil microorganisms (autoclaved or natural soil). Under the highest concentrations and autoclaved soil, more than 80 % of target species were germination and/or growth-inhibited, and only two species were non-sensitive. Under more natural conditions (lower extracts concentrations and natural soil with microorganisms), only 50 % of species were still inhibited, one was non-sensitive, and five were stimulated. Thus, microorganisms alter the expression of allelochemicals released into the ecosystem, which highlights their key role in chemical plant-plant interactions. The results of allelopathic experiments conducted in the lab are consistent with the community patterns observed in the field. These findings suggest that allelopathy is likely to shape vegetation composition and participate to the control of biodiversity in Mediterranean open mosaic habitats.