991 resultados para Soil communities


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Diffusion controls the gaseous transport process in soils when advective transport is almost null. Knowledge of the soil structure and pore connectivity are critical issues to understand and modelling soil aeration, sequestration or emission of greenhouse gasses, volatilization of volatile organic chemicals among other phenomena. In the last decades these issues increased our attention as scientist have realize that soil is one of the most complex materials on the earth, within which many biological, physical and chemical processes that support life and affect climate change take place. A quantitative and explicit characterization of soil structure is difficult because of the complexity of the pore space. This is the main reason why most theoretical approaches to soil porosity are idealizations to simplify this system. In this work, we proposed a more realistic attempt to capture the complexity of the system developing a model that considers the size and location of pores in order to relate them into a network. In the model we interpret porous soils as heterogeneous networks where pores are represented by nodes, characterized by their size and spatial location, and the links representing flows between them. In this work we perform an analysis of the community structure of porous media of soils represented as networks. For different real soils samples, modelled as heterogeneous complex networks, spatial communities of pores have been detected depending on the values of the parameters of the porous soil model used. These types of models are named as Heterogeneous Preferential Attachment (HPA). Developing an exhaustive analysis of the model, analytical solutions are obtained for the degree densities and degree distribution of the pore networks generated by the model in the thermodynamic limit and shown that the networks exhibit similar properties to those observed in other complex networks. With the aim to study in more detail topological properties of these networks, the presence of soil pore community structures is studied. The detection of communities of pores, as groups densely connected with only sparser connections between groups, could contribute to understand the mechanisms of the diffusion phenomena in soils.

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Soil is well recognized as a highly complex system. The interaction and coupled physical, chemical, and biological processes and phenomena occurring in the soil environment at different spatial and temporal scales are the main reasons for such complexity. There is a need for appropriate methodologies to characterize soil porous systems with an interdisciplinary character. Four different real soil samples, presenting different textures, have been modeled as heterogeneous complex networks, applying a model known as the heterogeneous preferential attachment. An analytical study of the degree distributions in the soil model shows a multiscaling behavior in the connectivity degrees, leaving an empirically testable signature of heterogeneity in the topology of soil pore networks. We also show that the power-law scaling in the degree distribution is a robust trait of the soil model. Last, the detection of spatial pore communities, as densely connected groups with only sparser connections between them, has been studied for the first time in these soil networks. Our results show that the presence of these communities depends on the parameter values used to construct the network. These findings could contribute to understanding the mechanisms of the diffusion phenomena in soils, such as gas and water diffusion, development and dynamics of microorganisms, among others.

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Debido al futuro incierto de la mayor parte de los fumigantes edáficos usados actualmente en la Unión Europea, que pueden implicar riesgos para la salud humana/animal y el medio ambiente, es necesario desarrollar programas de manejo integrado para el control de plagas de cultivos. Estos programas se incluyen como obligatorios en el Reglamento (EC) No. 1107/2009. De acuerdo con este Reglamento, es obligatoria la evaluación del riesgo asociado al uso de productos fitosanitarios sobre los organismos edáficos no diana y sus funciones, además de llevar a cabo ensayos con diferentes especies indicadoras para obtener datos de toxicidad que puedan ser usados posteriormente en la evaluación de riesgo. Sin embargo, la baja representatividad de algunas de estas especies indicadoras en el área Mediterránea supone una gran limitación. En esta situación, el Panel Científico de Productos Fitosanitarios y sus Residuos de la Autoridad Europea en Seguridad Alimentaria (EFSA), ha señalado la necesidad de modificar los datos ecotoxicológicos requeridos para evaluar los efectos adversos de los productos fitosanitarios de una manera más integrada, incluyendo criterios funcionales y estructurales mediante organismos como bacterias, hongos, protozoos y nematodos. De este modo, la EFSA ha recomendado el uso de los nematodos en la evaluación de la funcionalidad y estructura del suelo. Los nematodos están globalmente distribuidos y son morfológicamente diversos; esto junto con su gran abundancia y diversidad de respuestas a las perturbaciones edáficas, los convierte en indicadores adecuados del estado del suelo. Puesto que los nematodos interaccionan con muchos otros organismos que participan en diferentes eslabones de la red trófica edáfica, jugando papeles importantes en procesos edáficos esenciales en los agroescosistemas, la diversidad de nematodos es, a menudo, usada como indicador biológico de los efectos de las prácticas agrícolas en el estado del suelo. En los últimos años, diferentes índices basados en la comunidad nematológica han facilitado la interpretación de datos complejos sobre la ecología del suelo. Los índices de la red trófica edáfica, basados en la abundancia de grupos funcionales definidos como grupos C-P y grupos tróficos, permiten la evaluación de la funcionalidad de la red trófica edáfica. Por otra parte, la dificultad en la identificación taxonómica de nematodos para explicar su uso limitado como indicadores ecológicos, es ampliamente discutida, y existe cierta controversia en cuanto a la eficacia de los diferentes métodos de identificación de nematodos. Se argumenta que la identificación morfológica es difícil y puede llevar mucho tiempo debido a la falta de expertos especializados, y se afirma que las técnicas moleculares pueden resolver algunas limitaciones de las técnicas morfológicas como la identificación de juveniles. Sin embargo, los métodos de identificación molecular tienen también limitaciones; la mayoría de las bases de datos de secuencias de ADN están fuertemente orientadas hacia los nematodos fitoparásitos, los cuales representan sólo una parte de la comunidad edáfica de nematodos, mientras que hay poca información disponible de nematodos de vida libre a pesar de representar la mayoría de los nematodos edáficos. Este trabajo se centra en el estudio de los efectos de fumigantes edáficos en la funcionalidad del suelo a través del uso de diferentes indicadores basados en la comunidad de nematodos, como los índices de la red trófica, índices de diversidad, abundancia de los taxones más relevantes etc. También se han analizado otros indicadores funcionales relacionados con la supresividad edáfica, el ciclo de nutrientes o la actividad de la microfauna del suelo. En el capítulo 1, la diversidad de nematodos estudiada en una explotación comercial de fresa y sus alrededores durante dos campañas consecutivas en el suroeste español, fue baja en los suelos fumigados con fumigantes químicos ambas campañas y, aunque se observó una recuperación a lo largo de la campaña en la zona tratada, los suelos fumigados mostraron una condición perturbada permanente. La comunidad de nematodos estuvo más asociada al ciclo de nutrientes en la zona sin cultivar que en los suelos cultivados, y se observó poca relación entre la biomasa de las plantas y la estructura de la comunidad de nematodos. Los surcos sin tratar dentro de la zona de cultivo funcionaron como reservorio tanto de nematodos fitoparásitos como beneficiosos; sin embargo estas diferencias entre los surcos y los lomos de cultivo no fueron suficientes para mantener la supresividad edáfica en los surcos. Los suelos tratados fueron menos supresivos que los suelos sin tratar, y se observaron correlaciones positivas entre la supresividad edáfica y la estructura de la red trófica edáfica y la diversidad de nematodos. En el capítulo 2, se evaluaron los efectos de dos pesticidas orgánicos con efecto nematicida y dos nematicidas convencionales sobre las propiedades físico químicas del suelo, la diversidad de nematodos y la biomasa de las plantas en condiciones experimentales en dos tipos de suelo: suelos agrícolas poco diversos y suelos provenientes de una zona de vegetación natural muy diversos. El mayor efecto se observó en el tratamiento con neem, el cual indujo un gran incremento en el número de dauerlarvas en los suelos pobres en nutrientes, mientras que el mismo tratamiento indujo un incremento de poblaciones de nematodos bacterívoros, más estables y menos oportunistas, en los suelos del pinar ricos en materia orgánica. En el capítulo 3, se comparó la eficacia de métodos moleculares (TRFLP, Terminal Restriction Fragment Length Polymorphism) y morfológicos (microscopía de alta resolución) para la identificación de diferentes comunidades denematodos de España e Irlanda. Se compararon estadísticamente las diferencias y similitudes en la diversidad de nematodos, otros indicadores ecológicos y de la red trófica edáfica. Las identificaciones mediante el uso de TRFLP sólo detectó un porcentaje de los taxones presentes en las muestras de suelo identificadas morfológicamente, y los nematodos omnívoros y predadores no fueron detectados molecularmente en nuestro estudio. Los índices calculados en base a los nematodos micróboros mostraron más similitud cuando se identificaron morfológica y molecularmente que los índices basados en grupos tróficos más altos. Nuestros resultados muestran que, al menos con la técnica usada en este estudio, la identificación morfológica de nematodos es una herramienta fiable y más precisa que la identificación molecular, puesto que en general se obtiene una mayor resolución en la identificación de nematodos. En el capítulo 4, se estudiaron también los efectos de los nematicidas químicos sobre la comunidad de nematodos y la biomasa de las plantas en condiciones experimentales de campo, donde se aplicaron en una rotación de cultivo judía-col durante un ciclo de cultivo. Se aplicaron dos tipos de enmiendas orgánicas con el objetivo de mitigar el efecto negativo de los productos fitosanitarios sobre la diversidad edáfica. El efecto de los nematicidas sobre las propiedades del suelo y sobre la comunidad de nematodos fue más agudo que el efecto de las enmiendas. La incorporación de los restos de cosecha al final del ciclo de cultivo de la judía tuvo un gran efecto sobre la comunidad de nematodos, y aunque el número total de nematodos incrementó al final del experimento, se observó una condición perturbada permanente de la red trófica edáfica a lo largo del experimento. ABSTRACT Due to the uncertain future of the soil fumigants most commonly used in the EU, that might involve risks for human/animal health and the environment, there is a need to develop new integrated pest management programs, included as mandatory in the Regulation (EC) No. 1107/2009, to control crop diseases. According to this Regulation, evaluating the risk associated to the use of the plant production products (PPP) on non-target soil fauna and their function, and developing assays with different indicator species to obtain toxicity data to be used in the risk evaluation is mandatory. However, the low representativeness of some of these indicator species in the Mediterranean area is a relevant limitation. In this situation, the Scientific Panel of Plant Protection Products and their Residues of the European Food Safety Authority (EFSA) has pointed out the necessity of modifying the ecotoxicological data set required to evaluate non-target effects of PPP in a more integrated way, including structural and functional endpoints with organism such as bacteria, fungi, protists and nematodes. Thus, EFSA has recommended the use of nematodes in the assessment of the functional and structural features of the soil. Nematodes are globally distributed and morphologically diverse, and due to their high abundance and diversity of responses to soil disturbance, they are suitable indicators of the soil condition. Since nematodes interact with many other organisms as participants in several links of the soil food web, playing important roles in essential soil processes in agroecosystems, nematode diversity is often used as a biological indicator of the effects of agricultural practices on soil condition. In the last years, various indices based on soil nematode assemblages, have facilitated the interpretation of complex soil ecological data. Soil food web indices based on the abundances of functional guilds defined by C-P groups and trophic groups, permit evaluating soil food web functioning. On the other hand, the difficulty of nematode taxonomical identification is commonly argued to explain their limited used as ecological indicators, and there is a certain controversy in terms of the efficacy of various nematode identification methods. It is argued that the morphological identification is difficult and time consuming due to the lack of specialist knowledge, and it is claimed that molecular techniques can solve some limitations of morphological techniques such as the identification of juveniles. Nevertheless, molecular identification methods are limited too, since most of the DNA-based databases are strongly oriented towards plant-parasitic nematodes that represent only a fraction of the soil nematode community, while there is little information available on free-living nematodes, which represent most soil nematodes. This work focuses on the study of the effects of soil fumigants on soil functioning through the use of different indicators based on soil nematode community as soil food web indices, diversity indices, the abundance of more relevant taxa etc. Other functional indicators related to soil suppressiveness, nutrient cycling, or the activity of soil microfauna have been also studied. In chapter 1, nematode diversity assessed in a commercial strawberry farm and its surroundings for two consecutive growing seasons in southern Spain, was low in fumigated soils with chemical pesticides throughout both seasons and, although yearly recovery occurred within the treated fields, fumigated soils showed a permanent perturbed condition. The nematode community was more closely associated to nutrient cycling in the non-cropped than in the cropped soils, and the link between plant biomass and nematode community structure was weak. Non-treated furrows within the treated fields were a reservoir of both beneficial and plant-parasitic nematodes, but such difference between furrows and beds was not enough to maintain more suppressive soil assemblages in the furrows. Treated soils were less suppressive than unmanaged soils, and there was a positive and significant correlation between soil suppressiveness and soil food web structure and diversity. In chapter 2, the effects of two organic pesticides with nematicide effect and two chemical nematicides on soil physicalchemical properties, soil nematode diversity and plant biomass in experimental conditions were assessed in two types of soils: low diversity soils from an agricultural farm, and high diversity soils from a natural vegetation area. The larger effect was observed on the neem treatment, which induced a large boost of dauer juveniles in the nutrient-depleted soil, while the same treatment induced the increase of more stable, less opportunistic, populations of generalist bacterivore nematodes in the pine forest soil, rich in organic matter. In chapter 3, comparison of the efficiency of molecular (TRFLP, Terminal Restriction Fragment Length Polymorphism) and morphological (microscopy at high magnification) identification methods was carried out in different nematode communities from five sites of different land uses in Spain and Ireland. Differences and similarities on nematode diversity and other ecological and soil food web indices assessed by both methods, were statistically compared. Molecular identification with TRFLP only detected a percentage of the taxa present in the soil samples identified morphologically, and omnivores and predators were not detected molecularly in our study. Indices involving microbial feeding nematodes were more similar between identification methods than indices involving higher trophic links. Our results show that, at least with the technique used in this study, identifying nematodes morphologically is a reliable and more precise identification tool than molecular identification, since a higher taxonomic resolution is in general obtained compared to TRFLP. In chapter 4, the effect of chemical nematicides on nematode community descriptors and plant biomass was also studied in field conditions in an experimental area in which dazomet and dimethyl disulfide was applied in a bean-cabbage rotation system for a single season. Organic amendments were incorporated into the soil with the aim of mitigate the negative effect of the pesticides on soil diversity. The effect of the nematicides was much more noticeable than the effect of the amendments on soil properties and nematode community descriptors. The incorporation of bean crop residues into the soil at the end of bean crop cycle affected soil nematode community descriptors to a great extent, and although total number of nematodes increased at the end of the experiment, a permanent perturbed soil food web condition was observed along the experiment.

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Aim of study. Orchidaceae has the largest number of species of any family in the plant kingdom. This family is subject to a high risk of extinction in natural environments, such as natural parks and protected areas. Recent studies have shown the prevalence of many species of orchids to be linked to fungal soil diversity, due to their myco-heterotrophic behaviour. Plant communities determine fungal soil diversity, and both generate optimal conditions for orchid development. Area of study. The work was carried out in n the two most important natural parks in Alicante (Font Roja and Sierra Mariola), in South-eastern of Spain. Material and Methods. We designed a molecular tool to monitor the presence of Russula spp. in soil and orchids roots, combined with phytosociological methods. Main results. Using a PCR-based method, we detected the presence in the soil and Limodorum abortivum orchid roots of the mycorrhizal fungi Russula spp. The species with highest coverage was Quercus rotundifolia in areas where the orchid was present. Research highlights. We present a useful tool based on PCR to detect the presence of Russula spp. in a natural environment. These results are consistent with those obtained in different studies that linked the presence of the mycorrhizal fungi Russula spp. in roots of the species Limodorum and the interaction between these fungal species and Quercus ilex trees in Mediterranean forest environments.

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Large parts of the eastern half of the Tibetan Plateau are covered between (3,500) 4,000 and nearly 6,000 m a.s.l. by alpine sedge mats (key species Kobresia pygmea), which attain an extension of ca. 450,000 km**2. It is considered to be the world's largest alpine ecosystem. Moreover, there exist isolated (relic) forests in the same area up to an altitude of 4,700 m a.s.l. mainly consisting of juniper (Juniperus) and spruce (Picea). Large parts of the Kobresia ecosystem are expected to be a grazing-resistant replacement formation, replacing forests and grass-dominated plant communities due to human and/or climatic impact. Recently, a research project was launched to increase knowledge about the properties and genesis of these forests and sedge mats (Present-day dynamics and Holocene landscape history of fragmented forest biocoenoses in Tibet; headed by G. Miehe, Marburg).

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Mode of access: Internet.

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Thesis (Master's)--University of Washington, 2016-06

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A study of the prevalence, intensity and risk factors for soil-transmitted helminth infection was undertaken among school children aged 5-9 years attending a primary school in the fishing village in Peda Jalaripet, Visakhapatnam, South India. One hundred and eighty nine (92.6%) of 204 children were infected with one or more soil transmitted helminth parasites. The predominant parasite was Ascaris lumbricoides (prevalence of 91%), followed by Trichuris trichiura (72%) and hookworm (54%). Study of age-specific prevalence and intensity of infection revealed that the prevalence and intensity of A. lumbricoides infection was higher among younger children than older children. While aggregation of parasite infection was observed, hookworm infection was more highly aggregated than either A. lumbricoides or T. trichiura. Multivariate analysis identified parental occupation, child's age and mother's education as the potential risk factors contributing to the high intensity of A. lumbricoides infection. Children from fishing families with low levels of education of the mother had the highest intensity of A. lumbricoides infection. As the outcome of chemotherapy programs to control soil transmitted helminth infection is dependant on the dynamics of their transmission, there is a need for further studies to better define the role of specific factors that determine their prevalence, intensity and aggregation in different epidemiological settings. (C) 2004 Elsevier B.V. All rights reserved.

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A soil suspension was used as a source to initiate the development of microbial communities in flow cells irrigated with 2,4-dichlorophenoxyacetic acid (2,4-D) (25 mu g ml(-1)). Culturable bacterial members of the community were identified by 16S rRNA gene sequencing and found to be members of the genera Pseudomonas, Burkholderia, Collimonas and Rhodococcus. A 2,4-D degrading donor strain, Pseudomonas putida SM 1443 (pJP4::gfp), was inoculated into flow cell chambers containing 2-day old biofilm communities. Transfer of pJP4::gfp from the donor to the bacterial community was detectable as GFP fluorescing cells and images were captured using confocal scanning laser microscopy (GFP fluorescence was repressed in the donor due to the presence of a chromosomally located lacl(q) repressor gene). Approximately 5-10 transconjugant microcolonies, 20-40 mu m in diameter, could be seen to develop in each chamber. A 2,4-D degrading transconjugant strain was isolated from the flow cell system belonging to the genus Burkholderia.

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Biological soil crusts (BSCs) are formed by aggregates of soil particles and communities of microbial organisms and are common in all drylands. The role of BSCs on infiltration remains uncertain due to the lack of data on their role in affecting soil physical properties such as porosity and structure. Quantitative assessment of these properties is primarily hindered by the fragile nature of the crusts. Here we show how the use of a combination of non-destructive imaging X-ray microtomography (XMT) and Lattice Boltzmann method (LBM) enables quantification of key soil physical parameters and the modeling of water flow through BSCs samples from Kalahari Sands, Botswana. We quantify porosity and flow changes as a result of mechanical disturbance of such a fragile cyanobacteria-dominated crust. Results show significant variations in porosity between different types of crusts and how they affect the flow and that disturbance of a cyanobacteria-dominated crust results in the breakdown of larger pore spaces and reduces flow rates through the surface layer. We conclude that the XMT–LBM approach is well suited for study of fragile surface crust samples where physical and hydraulic properties cannot be easily quantified using conventional methods.

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Oribatid mites are one of the most abundant groups of the ground-dwelling mesofauna. They can be found in almost every terrestrial habitat all over the world and they are characterized by great species richness and great number of individuals. In spite of that not enough is known about their behaviour on community level and their spatial and temporal pattern in different habitats of the world. In our present study the seasonal behaviour of oribatid mite communities was analysed in three types of microhabitats in a temperate deciduous forest: in leaf litter, soil and moss. Samples were collected at a given site in a year and a half and the oribatid mite communities living there were studied on genus level along with the changes of meteorological factors characteristic of the area. The results show that corresponding to similar previous researches, the communities in our study do not have a seasonally changing, returning pattern either. Based on this, we can conclude that climatic differences and differences in other seasonally changing factors between the seasons do not have a significant role in the annual change of communities. Besides that we discovered that the communities of the three microhabitats are not completely the same. It is the oribatid mite community of the moss which differs mostly from communities in the leaf litter and in the soil. Our study calls attention among others to the fact that compositional changes of the oribatid mite communities living all over the world and their causes are unclear to date.

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Hydroperiod and nutrient status are known to influence aquatic communities in wetlands, but their joint effects are not well explored. I sampled floating periphyton mat and flocculent detritus (floc) infaunal communities using 6-cm diameter cores at short- and long-hydroperiod and constantly inundated sites across a range of phosphorus (P) availability (total phosphorus in soil, floc and periphyton). Differences in community structure between periphyton and floc microhabitats were greater than any variation attributable to hydroperiod, P availability, or other spatial factors. Multivariate analyses indicated community structure of benthic-floc infauna was driven by hydroperiod, although crowding (no. g−1 AFDM) of individual taxa showed no consistent responses to hydroperiod or P availability. In contrast, community structure of periphyton mat infauna was driven by P availability, while densities of mat infauna (no. m−2) were most influenced by hydroperiod (+correlations). Crowding of mat infauna increased significantly with P availability in short-hydroperiod marshes, but was constant across the P gradient in long-hydroperiod marshes. Increased abundance of floating-periphyton mat infauna with P availability at short-hydroperiod sites may result from a release from predation by small fish. Community structure and density were not different between long-hydroperiod and constantly inundated sites. These results have implications for the use of macroinvertebrates as indicators of water quality in wetlands and suggest the substrate sampled can influence interpretation of ecological responses observed in these communities.

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Tree islands in the Everglades wetlands are centers of biodiversity and targets of restoration, yet little is known about the pattern of water source utilization by the constituent woody plant communities: upland hammocks and flooded swamp forests. Two potential water sources exist: (1) entrapped rainwater in the vadose zone of the organic soil (referred to as upland soil water), that becomes enriched in phosphorus, and (2) phosphorus-poor groundwater/surface water (referred to as regional water). Using natural stable isotope abundance as a tracer, we observed that hammock plants used upland soil water in the wet season and shifted to regional water uptake in the dry season, while swamp forest plants used regional water throughout the year. Consistent with the previously observed phosphorus concentrations of the two water sources, hammock plants had a greater annual mean foliar phosphorus concentration over swamp forest plants, thereby supporting the idea that tree island hammocks are islands of high phosphorus concentrations in the oligotrophic Everglades. Foliar nitrogen levels in swamp forest plants were higher than those of hammock plants. Linking water sources with foliar nutrient concentrations can indicate nutrient sources and periods of nutrient uptake, thereby linking hydrology with the nutrient regimes of different plant communities in wetland ecosystems. Our results are consistent with the hypotheses that (1) over long periods, upland tree island communities incrementally increase their nutrient concentration by incorporating marsh nutrients through transpiration seasonally, and (2) small differences in micro-topography in a wetland ecosystem can lead to large differences in water and nutrient cycles.

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Belowground biomass is a critical factor regulating ecosystem functions of coastal marshes, including soil organic matter (SOM) accumulation and the ability of these systems to keep pace with sea-level rise. Nevertheless, belowground biomass responses to environmental and vegetation changes have been given little emphasis marsh studies. Here we present a method using stable carbon isotopes and color to identify root and rhizomes of Schoenoplectus americanus (Pers.) Volk. ex Schinz and R. Keller (C3) and Spartina patens (Ait.) Muhl. (C4) occurring in C3− and C4-dominated communities in a Chesapeake Bay brackish marsh. The functional significance of the biomass classes we identified is underscored by differences in their chemistry, depth profiles, and variation in biomass and profiles relative to abiotic and biotic factors. C3 rhizomes had the lowest concentrations of cellulose (29.19%) and lignin (14.43%) and the lowest C:N (46.97) and lignin:N (0.16) ratios. We distinguished two types of C3 roots, and of these, the dark red C3 roots had anomalously high C:N (195.35) and lignin:N (1.14) ratios, compared with other root and rhizome classes examined here and with previously published values. The C4-dominated community had significantly greater belowground biomass (4119.1 g m−2) than the C3-dominated community (3256.9 g m−2), due to greater total root biomass and a 3.6-fold higher C3-root:rhizome ratio in the C4-dominated community. C3 rhizomes were distributed significantly shallower in the C4-dominated community, while C3 roots were significantly deeper. Variability in C3 rhizome depth distributions was explained primarily by C4 biomass, and C3 roots were explained primarily by water table height. Our results suggest that belowground biomass in this system is sensitive to slight variations in water table height (across an 8 cm range), and that the reduced overlap between C3 and C4 root profiles in the C4-dominated community may account for the greater total root biomass observed in that community. Given that future elevated atmospheric CO2 and accelerated sea-level rise are likely to increase C3 abundance in Atlantic and Gulf coast marshes, investigations that quantify how patterns of C3 and C4 belowground biomass respond to environmental and biological factors stand to improve our understanding of ecosystem-wide impacts of global changes on coastal wetlands.