Multiple stages of plant root calcification deciphered by chemical and micromorphological analyses

Rhizoliths, that is, roots fossilized by secondary carbonates, have been known for ages and are increasingly used for paleoenvironmental reconstructions. However, knowledge about their formation mechanisms remains limited. This study reports the mineralogical and chemical characterization of rhizoliths at different stages of mineralization and fossilization in the Late Pleistocene loess–paleosol sequence of Nussloch (SW Germany). Scanning electron microscopy coupled with elemental mapping and ¹³C solid‐state nuclear magnetic resonance were used to concomitantly characterize the mineral and organic matter of the rhizoliths. These joint analyses showed for the first time that large rhizoliths are not necessarily remains of single large roots but consist of numerous microrhizoliths as remains of fine roots, formed mainly by calcium carbonates with only low amounts of Mg and Si. They further revealed that the precipitation of secondary carbonates occurs not only around, but also within the plant root and that fossilization leads to the selective preservation of recalcitrant root biopolymers—lignin and suberin. The precipitation of secondary carbonates was observed to occur first around fine roots, the epidermis acting as a first barrier, and then within the root, within the cortex cells, and even sometimes around the phloem and within the xylem. This study suggests that the calcification of plant roots starts during the lifetime of the plant and continues after its death. This has to be systematically investigated to understand the stratigraphic context before using (micro)rhizoliths for paleoenvironmental reconstructions in terrestrial sediments.     

Can orchid mycorrhizal fungi be persistently harbored by the plant host?

The environmental distribution of non-obligate orchid mycorrhizal (OM) symbionts belonging to the ‘rhizoctonia’ complex remains elusive. Some of these fungi, indeed, are undetectable in soil outside the host rhizosphere. A manipulation experiment was performed to assess the importance of neighbouring non-orchid plants and soil as possible reservoirs of OM fungi for Spiranthes spiralis, a widespread photosynthetic European terrestrial orchid species. Fungi of S. spiralis roots were identified by DNA metabarcoding before and 4 months after the removal of the surrounding vegetation and soil. Although such a treatment significantly affected fungal colonization of newly-formed orchid roots, most OM fungi were consistently associated with the host roots. Frequency patterns in differently aged roots suggest that these fungi colonize new orchid roots from either older roots or other parts of the same plant, which may thus represent an environmental source for the subsequent establishment of the OM symbiosis.     

西沙群岛石岛根管石丛的形成分析*

根管石是高等植物的化石, 它不仅能证实古土壤的存在, 还是古植被重建、古气候反演和沉积环境重建的良好载体。根管石在中国南海西沙群岛石岛东西海岸的潮上带广泛发育, 而在南海其他岛屿鲜有报道。石岛的绝大部分根管石在岩壁及古土壤层中独立分布, 此外还发育了裸露的根管石丛, 形态保存相对完好。本文运用X-射线断层扫描成像技术(Micro-CT)针对石岛根管石丛进行三维可视化分析, 获取其精细的形态特征, 结合植物生态学特性和根管石丛的微观结构, 探讨了西沙群岛石岛根管石丛的形成, 以此为海岸带环境下根管石的形成机制和沉积环境重建提供科学依据。结果表明: X-射线断层扫描得到的石岛根管石丛宏观三维形貌更加直观地展示三层环带状构造。结合岩石薄片, 推测石岛根管石丛内带和中带皆由植物自身交代形成, 而外带则为珊瑚砂固结成岩。植被的形态特征和生态适应性表明石岛根管石丛由茎干形成, 建议使用“根茎石”这一术语更为贴切。同时, 初步推测其原生植被是分布于石岛的优势种草海桐。     

自然降雨下黄土丘陵区草灌植物垂直覆盖结构的减流减沙效应

黄土高原是我国水土流失和生态环境问题最为严重的地区之一,植被恢复是防治水土流失的重要措施。植物垂直覆盖结构包括地上冠层、地表枯落物和地下根系,各组分具有不同的水土保持作用,是研究植被与水土流失关系的基本单元。目前,关于植物垂直覆盖结构不同组分对土壤侵蚀影响的研究主要是基于人工模拟降雨,缺少自然降雨条件下不同植物的垂直覆盖结构对产流、产沙和入渗等多过程影响的系统研究。本研究以黄土丘陵区典型的草本(须芒草)、半灌木(铁杆蒿)和灌木(绣线菊)为研究对象,每种植物进行三种处理(自然状态、去除枯落物和仅留根系)以及裸地对照,观测2015—2016年降雨事件的产流产沙量和入渗量,分析植物不同垂直覆盖结构的减流减沙效益及其相对贡献。结果表明:三种植物均具有较好的减流(45.9%—73.2%)、减沙效益(87.5%—94.6%)和增加入渗作用(4.7%—10.8%),灌木的减流效果(73.2%)显著高于草本(45.9%)和半灌木(63.5%),但三种植物间的减沙效益没有显著性差异。冠层的减流作用最大,贡献率接近一半(48%—50%),草本枯落物的减流贡献率与根系基本一致,而半灌木和灌木枯落物的减流贡献...     

A molecular marker-based assessment of sedimentary organic matter sources and distributions in Florida Bay

Seven surface sediment samples covering the general geographical area of Florida Bay were examined through the measurement of ¹³C isotopic abundance and lipid classes to assess the distributions and sources of organic matter (OM) in this estuarine environment. The bulk δ¹³C_org value shifted from a more isotopically depleted (−19.9‰) to a more isotopically enriched (−13.5‰) signal along the NE to SW transect. Two geochemical proxies (Paq and C₂₅/C₂₇ n-alkan–2-ones) indicative of seagrass-derived OM significantly increased from near-shore to offshore areas, while taraxerol, a biomarker for mangroves, substantially decreased from 7200 to 284 ng/g along that transect. A clear spatial variation of OM sources was observed in Florida Bay. Generally, the sites in the NE contained mixed OM sources of terrestrial (mangrove) and seagrass-derived OM, where the terrestrial component accounted for over 60% of the OM. In contrast, the sites in central and SW Florida Bay were strongly dominated by seagrass-derived OM. Other lipid fractions such as fatty acids, n-alcohols and sterols revealed an important contribution of algae and bacteria especially in the central and SW section of the Bay. Relatively abundant C₂₅ HBIs suggest important marine diatom inputs, while the presence of C₂₀ HBIs particularly in central Florida Bay possibly reflects the contribution of cyanobacterial mats. The molecular proxies developed in this study to assess OM sources in Florida Bay are promising tools for the characterization and seasonal variability assessment of OM in this and other similar subtropical and tropical estuaries and for paleoenvironmental studies. Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users.     

Determining the onset of autumn grass senescence in subtropical sour-veld grasslands using remote sensing proxies and the breakpoint approach

Information on the onset of autumn grass senescence in subtropical grasslands is essential for ascertaining the duration of poor forage quality. It is well-established that during senescence, grass leaves lose their nutrients to the rooting systems; which affects the quality and quantity of forage resources. The timing of the onset of autumn grass senescence is critical in determining the potential lifespan through which the provision of quality forage can be sustained in a grazing area. However, objective and robust methods for estimating the onset of autumn grass senescence at a rangeland scale are limited. Hence, this study sought to characterize the onset of autumn senescence in mesic subtropical sour-veld grasslands using remotely sensed data. Ten monthly vegetation indices were generated from the Sentinel 2 data and used as proxies to explain the onset of autumn grass senescence. The performance of the proxies was validated using the corresponding field-measured monthly grass chlorophylls. Results showed that the Chlorophyll Red-Edge (CHL-RED-EDGE) and the Normalized Difference Red Edge Index (NDVI₇₀₅) were the most important proxies for characterizing the autumn grass senescence. In addition, monthly (i.e., January to June) mean values of the two best proxies were fitted in a piecewise linear regression model with a breakpoint approach to determine the start of the autumn grass senescence. The first proxy (i.e., NDVI₇₀₅) predicted that the grass in the study area starts senescing on day number ± 98 of the year (R² = 0.97, RMSE = 0.024) while the second (i.e., CHL-RED-EDGE) suggested day number ± 106 of the year (R² = 0.96, RMSE = 0.052). Overall, this study demonstrated the value of remote sensing proxies in estimating the autumn grass senescence and in determining its onset. These results provide a basis for understanding the impact of autumn senescence on foraging resource provision in rangeland ecosystems.     

Impact of Anthropogenic Organic Matter on the Distribution Patterns of Sediment Microbial Community from the Yangtze River,China

Abstract

How microbes respond to substantial and increasing anthropogenic disturbance remains an open question in river systems. We tested the hypothesis that the source and distribution of anthropogenic organic matter (OM) were significant factors affecting the spatial variation of the microbial community composition of the Yangtze River sediments. Bulk geochemical proxies and lignin phenols suggested a general decrease of terrestrial C3 plants or soil OM input from the middle to the lower reaches. Fecal sterols inferred higher sewage contamination levels in the middle reaches. Polycyclic aromatic hydrocarbons (PAHs) distribution indicated a dominant biomass and coal combustion signal in the middle reaches, whereas a mixed source including petroleum combustion in the lower reaches. Phylogenetic analysis revealed a large portion of Methanobacteria and Verrucomicrobia enriched in the middle reaches, whereas OM-degrading bacteria, including Flavobacteria, Acidobacteria, and Alphaproteobacteria were dominant in the lower reaches. Quantitative PCR analyses and multivariate analysis further demonstrated that sources and distribution of OM had combined effects in shaping alpha and beta-diversity of sediment microbial communities. Sewage discharge and incomplete OM combustion, respectively, were associated with Methylococcaceae, Chloroflexi, and Bacteroidetes groups. This study provides a foundation for further understanding of the river sediment microbial composition, considering the continued increase of anthropogenic influences.     

黄土区露天煤矿排土场不同微地貌土壤种子库特征

黄土区露天煤矿排土场边坡水力冲刷强烈,不同植被恢复模式与水力作用耦合形成的不同微地貌可通过对种子的不同拦截作用而影响植被的恢复及发展。为了探讨排土场不同微地貌对种子的拦截效益及不同植被恢复模式的恢复潜力,选取不同恢复模式下形成的细沟、丛岛和草带三种微地貌为研究对象,以邻近裸坡为对照,研究不同微地貌土壤种子库特征及其拦截力。结果表明：（1）在不同微地貌中土壤种子库均以一年生藜科和禾本科植物为主;丛岛微生境的土壤种子库中有较丰富的物种和较高的多样性指数。（2）各微地貌的土壤种子库均具有表层聚集现象;草带微地貌中的土壤种子库总储量最大,且草带的0-2 cm土层处的土壤种子库密度显著高于各微地貌的各个土层（P<0.05）。（3）细沟和丛岛在上坡位对种子的拦截效果更好,草带在下坡位对种子的拦截效果更好。综上所述,沙蒿恢复模式形成的丛岛有利于增加土壤种子库的物种多样性,草带有利于增加土壤种子库的储量,故在黄土区排土场进行植被恢复时可以利用不同微地貌对土壤种子库分布特征的优势构建适宜的复合植被恢复模式。     

Environmental perspectives of Vetiveria zizanioides (L.) Nash

The twentieth century witnessed indiscriminate usage of natural resources for energy generation and xenobiotic chemical compounds for sustainability in agriculture and infrastructural development. Heavy metal and non-degradable chemical contamination of soil and water is one of the major environmental threats. In recent years, worldwide researchers are concentrating on the exploration of various sustainable methods to mitigate such environmental contamination. Vetiver (Vetiveria zizanioides (L.) Nash), a grass, is a proven source to mitigate such pollution, and in present days is one of the most recent thrust areas for the purpose of environmental mitigation. Unique morphology, physiology and symbiotic association render vetiver capable of tolerating environmental extremities. In addition, vetiver is also helpful in degradation of most of the recalcitrant compounds such as benzo[a]pyrene. The present review reflects the environmental perspectives of vetiver grass, a potential field which led the World Bank to initiate vetiver grass technology (VGT), which is now known as vetiver system (VS), in India and most of the other Asian countries to restore the natural environmental conditions.     

Radiostrontium contamination of soil and vegetation within the Semipalatinsk test site

The Semipalatinsk nuclear test site (STS) in the Republic of Kazakhstan was an important site for testing atomic bombs and other civil and military nuclear devices of the former Soviet Union. Results are presented from investigations on the extent of radiostrontium contamination in soils and vegetation at the technical areas of the STS, where the tests were conducted and in pastures used by farmers for grazing animals or for hay production. Our data are compared with those reported largely in the recent Russian language literature that has been reviewed. The extent of ⁹⁰Sr contamination of soil is highly variable over the STS with the highest values associated with the technical areas, particularly the Degelen mountains. Recently measured values in both the present data and the Russian language literature confirm the relatively high current contamination of soil and vegetation in the vicinity of tunnels and associated watercourses in the Degelen area. The proportion of ⁹⁰Sr in soil which could not be extracted with 6 M HCl was only an average of 20%, which is low compared to other test site areas and possibly indicates a relatively high mobility in this area, because the ⁹⁰Sr is derived from leakage from explosion tunnels along watercourses rather than being associated with fused silicates. A comparison of relative activity concentrations in soil and vegetation suggests that the transfer of ⁹⁰Sr to vegetation on the STS is high compared to that of ¹³⁷Cs and plutonium.     

Calcium Nodules as a Proxy for Quaternary Paleoclimate Change on China’s Loess Plateau

Different proxies have been used to investigate Quaternary paleoclimate change. Here, we used weathering of calcium nodules in paleosols on China’s Loess Plateau as a proxy for Quaternary paleoclimate changes to provide an alternative indicator of these changes. Paleosol and carbonate nodules were collected from Luochuan and Lantian counties in Shaanxi Province, China. We found that this approach allowed quantitative reconstruction of temperature, rainfall, soil mineral composition, and the effects of weathering and leaching. The changes in carbonate content in the loess and paleosol sequences were controlled by alternating dry and wet climatic conditions. Nodule formation conditions were directly affected by the leaching and migration of elements. The loess and paleosol sequences developed calcium nodules, and their formation was closely related to the rainfall and leaching characteristics of the paleoclimate. The paleoclimate and soil minerals affected the vegetation types and directly influenced changes in the soil. During formation of the calcium nodules, the surface vegetation evolved slowly, and the number of species and quantity of vegetation both decreased.     

模拟降雨下黄土坡面水沙过程对3种灌草植被垂直结构变化的响应

植被对坡面产流产沙过程的影响随植被类型及其垂直结构组分的变化而变化,然而这些因素如何影响坡面水沙过程却缺乏定量分析。利用野外径流小区和人工模拟降雨试验,研究了黄土丘陵区3种典型灌草（沙棘、柠条、苜蓿）及其不同垂直结构组分（叶、茎、枯落物、根系）对坡面产流和产沙过程的影响。结果表明：3种灌草均具有较好的减流减沙效益,且减沙作用强于减流作用,与裸地相比,灌草植被减少径流量32.49%-44.86%,减少侵蚀量72.99%-80.63%,降低坡面流速29.17%-45.83%。苜蓿的减流效益最佳,为44.86%,柠条的减沙效益最佳,为80.63%。3种植被的减流效益在不同产流时期差异明显,从产流初期到中期和后期逐渐减少,减沙效益在不同产流时期则没有明显变化。植被垂直结构不同组分对于减流减沙效益的相对贡献与其形态特征以及其空间分布方式有着密切的关系。地上部分对于减流效益和减速效益有较大的相对贡献率,平均为75.42%和68.38%,而不同植被茎、叶和枯落物的相对贡献具有一定的差异。根系则发挥较大的减沙作用,平均相对贡献为78.44%。植被垂直结构组分越完整,减流减沙效益越显著。研究对黄土丘陵区水土保持、植被恢复和建设提供重要的科学依据和理论指导。     

The ligament of Marshall as a parasympathetic conduit

The objective of the study was to investigate the morphology, distribution, and electrophysiological profile of the autonomic fibers that innervate the ligament of Marshall (LOM). Gross anatomical dissections were performed in 10 dogs. Sections of the left vagus nerve, left stellate ganglion, and the LOM were immunostained to identify adrenergic and cholinergic nerves. Hearts were also stained for acetylcholinesterase to identify epicardial cholinergic nerves. In vivo electrophysiological studies were performed in another 10 dogs before and after LOM ablation. The anatomical examination revealed that the LOM is innervated by a branch of the left vagus. Immunohistochemistry confirmed that these nerve bundles are predominantly cholinergic (cholinergic-to-adrenergic ratio of 12.6 +/- 3.9:1). Cholinergic nerves originating in the LOM were found to innervate surrounding left atrial structures, including the pulmonary veins, left atrial appendage, coronary sinus, and posterior left atrial fat pad. Ablation of the LOM significantly attenuated effective refractory period shortening at distant sites, such as pulmonary veins and left atrial appendage, in response to vagal stimulation (vagal-induced ERP decrease in the left atrium: baseline vs. postablation = 17 vs. 4%; P = 0.0056). In conclusion, the LOM contains a predominance of cholinergic nerve fibers. Cholinergic fibers arising from the LOM innervate surrounding structures and contribute to the electrophysiological profile of the left atrium. These findings may provide a basis for the role of the LOM in the genesis and maintenance of atrial fibrillation.     

Climate-related changes during the Late Glacial and early Holocene in northern Poland, as derived from the sediments of Lake Sierzywk

Reconstruction of past climate change and ecosystem response is important to correctly assess the impacts of global warming. In this study, we provide a paleoenvironmental record of in-lake and catchment changes in northern Poland during the Late Glacial and early Holocene using various biotic proxies (pollen, macrofossils and Cladocera) preserved in the lake sedimentary record. Chronology was derived from palynological correlation with a well-dated pollen sequence from nearby-lying Lake Ostrowite and some well-dated events of vegetation history in Central Europe. Pollen analysis provided information on regional climate change affecting vegetation dynamics, whereas macrofossils supplied substantial information on the response of local flora and fauna to climatic, geomorphological and limnological changes. Data were supplemented by analysis of Cladocera remains, which are of special importance because of their quick response to changes in trophic conditions and climate (especially temperature). The bottom of the sediment core reflects an initial stage of the lake formed during the late Aller?d. The Younger Dryas cooling apparently resulted in forest recession and presence of cold tolerant Cladocera species. Due to amelioration of climate at the end of the Younger Dryas and melting of ice, the lake deepened. The beginning of the Holocene was characterised by forest shrinkage and induced clear changes in local flora and fauna communities. The regional vegetation development deduced from the lake’s core is generally consistent with the vegetation history of central Europe. Due to the location of the site near the seashore (oceanic climate and western wind), signals of warming came earlier than inland and in eastern Poland.     

Recovery of native grass biodiversity by sowing on former croplands: Is weed suppression a feasible goal for grassland restoration?

Grassland restoration on former croplands offers good opportunity to mitigate the loss of grassland biodiversity. Weed suppression can be another benefit, which becomes increasingly important because of the high recent rate of abandonment of arable lands in Central and Eastern Europe. Our aim was to evaluate the usefulness of sowing two low-diversity seed mixtures followed by annual mowing, a frequently used restoration technique, in weed suppression. We found that rapidly forming cover of sown grasses effectively suppressed short-lived weeds and their germination except in the first year. The detected dense seed bank of short-lived weeds points out the possibility and threat of later weed infestation. In the short run perennial weeds cannot be suppressed easily by sowing and annual mowing. We found that the effectiveness of seed sowing followed by mowing in weed suppression can be different on sites with different history or seed mixture. Rapidly establishing perennial weeds, such as Agropyron species were only detected in former alfalfa fields; Cirsium arvense was found in former cereal and sunflower fields but not in former alfalfa fields. We found that the rate of weed suppression and success was influenced by the seed mixtures used. In several alkali restorations the high proportion of perennial weeds was detected in year 3. In loess restorations, much lower scores were typical. This was likely caused by the different seed mixture used. The loess seed mixture contained seeds of a clonally spreading tall-grass, Bromus inermis, which could compete more effectively with clonally spreading weeds, than could short grass species with or without tussock forming. Our findings indicate that post-restoration management require carefully designed actions that are fine-tuned addressing specific threats at the site level.     

Tree and grass rooting patterns in an African humid savanna

Abstract. Spatial and temporal soil partitioning between roots of the two savanna plant components, i.e. trees and grasses, were investigated in a West African humid savanna. Vertical root phytomass distribution was described for grass roots, large (> 2 mm) and fine (< 2 mm) tree roots, in open sites and beneath tree canopies. These profiles were established monthly over one year of vegetation growth. Natural ¹³C abundance measurement was used to determine the woody/herbaceous phytomass ratio in root samples. Tree and grass root distributions widely overlapped and both were mostly located in the top 20 cm of the soil. Grass root phytomass decreased with depth whereas woody root phytomass peaked at about 10 cm depth. No time partitioning was detected. These structural results do not support the hypothesis of soil resource partitioning between trees and grasses and are thus consistent with functional results previously reported.     

The emergence and intensification of early hunter‐gatherer niche construction

Hunter‐gatherers, especially Pleistocene examples, are not well‐represented in archeological studies of niche construction. However, as the role of humans in shaping environments over long time scales becomes increasingly apparent, it is critical to develop archeological proxies and testable hypotheses about early hunter‐gatherer impacts. Modern foragers engage in niche constructive behaviors aimed at maintaining or increasing the productivity of their environments, and these may have had significant ecological consequences over later human evolution. In some cases, they may also represent behaviors unique to modern Homo sapiens. Archeological and paleoenvironmental data show that African hunter‐gatherers were niche constructors in diverse environments, which have legacies in how ecosystems function today. These can be conceptualized as behaviorally mediated trophic cascades, and tested using archeological and paleoenvironmental proxies. Thus, large‐scale niche construction behavior is possible to identify at deeper time scales, and may be key to understanding the emergence of modern humans.     

Vesicular-arbuscular mycorrhizal inoculum potential affects the growth of Stryphnodendron microstachyum seedlings in a Costa Rican human tropical lowland

This study used a plant bioassay to investigate the vesicular-arbuscular mycorrhizal (VAM) inoculum potential of soil from three vegetation types (fern, secondary forest, and grass) in an abandoned pasture in the tropical humid lowlands at La Selva, in northeastern Costa Rica. Growth, measured as seedling height, number of leaves, and total (above- and belowground) biomass, of Stryphnodendron microstachyum Poepp. et Endl. (Synon. S. excelsum Harms) seedlings was significantly lower when grown in soil inoculum from the fern areas than in soil inoculum from the forest and grass areas. However, S. microstachyum seedlings grown in the fern inoculum had significantly greater VAM colonization than seedlings grown in the forest and grass inoculum. In addition, roots collected from a dominant plant species from each of the three vegetation types showed that the fern (Nephrolepsis biserrata) had significantly greater mycorrhizal colonization than the tree (Pentaclethra macroloba (Willd.) Kuntze or the grass (Brachiaria spp.). The results of this study suggest that differences in mycorrhizal inoculum potential among vegetation types and its effects on seedling growth may have important implications for the restoration and management of degraded lands.     

The effects of reed canary grass (<Emphasis Type="Italic">Phalaris arundinacea</Emphasis> L.) on wetland habitat and arthropod community composition in an urban freshwater wetland

Reed canary grass (Phalaris arundinacea L.) is an aggressive invader that dominates wetlands throughout the US. We examined the effects of reed canary grass on wetland habitat, both vegetation canopy architecture and soil environment, and its impacts the arthropod community in an urban wetland in Portland, OR, USA. Reed canary grass dominance resulted in reduced vegetation canopy complexity through reductions in native vegetation diversity and canopy height. In addition, reed canary grass dominance significantly changed the wetland soil environment, decreasing soil organic content and increasing soil moisture. The arthropod community responded to these habitat changes, being distinct between plots dominated by reed canary grass and those dominated by native vegetation. In addition, diversity measures were significantly lower in plots dominated by reed canary grass. Variables describing both vegetation canopy complexity and soil environment were more important predictors than relative abundance of reed canary grass in multiple regression models developed for dominant arthropod taxa and community metrics. Our results suggest that the mechanism by which reed canary grass affects the wetland arthropod community is primarily indirect, through habitat changes, rather than by directly altering its food source.     

Rainfall seasonality determines annual/perennial grass balance in vegetation of Mediterranean Iberian

Much recent attention has been focused on the invasion and dominance of annual grass species in areas thought to have been historically dominated by perennial life forms. Explanations of this phenomenon in the literature have focused on two mechanisms favoring annuals: ruderal strategy associated with disturbance, and stress escaping associated with dry sites or deserts. Here I present evidence from vegetation surveys at 50 sites across a 1,200 km band of the Iberian Peninsula—a source region for many invasive annuals—showing that relative annual versus perennial grass composition is not well correlated with degree of disturbance or average annual precipitation. However, annual dominance is strongly and significantly linked to the seasonality of precipitation, in particular the relative intensity of summer drought. Disturbance was significantly associated with annual grass dominance in Iberia, but with much less explanatory power than summer drought intensity. Slope, aspect, and soil parent material were not significantly correlated with annual versus perennial grass dominance. These results suggest that subtle differences in rainfall seasonality largely drive grass composition in herbaceous Mediterranean vegetation. Furthermore, the patterns of annual grass invasion observed in the world’s other Mediterranean climate regions may be associated with similar climatic drivers.