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Meiobenthos

Meiobenthos comprises the community of small benthic metazoans and foraminiferans, typically ranging in size from 32-64 μm to 1 mm, that inhabit the interstitial spaces within aquatic sediments and are operationally defined as organisms retained on sieves with mesh sizes between approximately 32 μm and 500-1000 μm.^[1]^[2]^[3] These organisms, often referred to as meiofauna in benthic contexts, bridge the size gap between microbenthos and macrobenthos, occupying niches in marine, estuarine, and freshwater environments worldwide.^[4] The composition of meiobenthos is dominated by nematodes, which often constitute over 80% of the total abundance, followed by harpacticoid copepods as the subdominant group, with additional contributions from annelids, tardigrades, rotifers, and gastrotrichs.^[5]^[6] This assemblage exhibits high taxonomic diversity, with species richness varying by habitat; for instance, in seagrass beds and intertidal zones, diversity is influenced by factors such as sediment granulometry and organic content.^[7]^[8] Meiobenthos communities are particularly abundant in shallow coastal sediments, but they also thrive in deep-sea environments, where they adapt to low oxygen and high-pressure conditions.^[9] Ecologically, meiobenthos plays a pivotal role in sediment processes, including the remineralization of organic matter through grazing and bioturbation, which enhances nutrient recycling and supports primary productivity in overlying waters.^[1] They serve as a critical trophic link, transferring energy from detritus and microalgae to higher-level consumers such as macrofauna, juvenile fish, and birds, thereby maintaining food web stability.^[8]^[10] Due to their sensitivity to environmental disturbances like pollution and hypoxia, meiobenthos assemblages are valuable indicators of ecosystem health, with shifts in diversity and abundance reflecting changes in habitat quality.^[10]^[11]

Definition and Characteristics

Definition and Size Criteria

Meiobenthos refers to a diverse assemblage of small, bottom-dwelling invertebrates and associated meiofauna that inhabit aquatic sediments, including marine, estuarine, and freshwater environments, and are distinguished from plankton, which drift passively in the water column, and nekton, which are active swimmers capable of directed locomotion.^[12] These organisms are integral to benthic communities, residing within interstitial spaces of sediments rather than the overlying water.^[4] The classification of meiobenthos is operational and size-based, typically encompassing metazoans and certain protozoans that are retained on sieves with a mesh size of 44–45 μm but pass through sieves with a mesh size of 500–1000 μm.^[3]^[13] This range excludes larger macrofauna, such as polychaetes or mollusks exceeding 1 mm, and smaller microbiota like bacteria or unicellular algae below 40 μm.^[14] While this sieve-based criterion provides a practical standard for sampling and study, exact thresholds vary across research protocols, with some studies using a lower limit of 40 μm or an upper limit of 1 mm to account for regional or habitat-specific adaptations.^[15] Metazoans, including nematodes and copepods, form the core, but protozoans such as foraminifers and ciliates are frequently included when they fall within the size range, contributing to the group's ecological dynamics.^[4] In marine sediments, meiobenthos exhibits high abundances, often ranging from 10³ to 10⁶ individuals per square meter, reflecting their adaptation to dense, interstitial lifestyles.^[4]^[16] Despite their minute size, their collective biomass can be comparable to that of macrofauna in certain coastal and sublittoral habitats, underscoring their significant role in sediment energy flow.^[17] Meiobenthic organisms are characterized by life history traits that enable rapid population responses to environmental changes, including short generation times of days to weeks and high reproductive rates.^[18] Parthenogenetic reproduction, where females produce offspring without fertilization, occurs in some groups like certain nematodes and rotifers, enhancing their resilience in variable sediment conditions.^[19]

Distinction from Other Benthic Groups

Meiobenthos are differentiated from microfauna, primarily composed of protozoans and other small eukaryotes smaller than approximately 44–100 μm, by their larger body sizes (typically 45–1000 μm), enhanced mobility that allows them to traverse interstitial pore spaces via water films, and a greater prevalence of predatory or grazing behaviors rather than being primarily decomposers; microfauna often engage in bacterivory or predation on microbes in a more localized manner within sediment matrices.^[20] In contrast to macrobenthos, which encompass larger invertebrates such as polychaetes and bivalves exceeding 500–1000 μm, meiobenthos occupy narrower interstitial voids in sediments, achieve substantially higher population densities (often orders of magnitude greater), and display faster metabolic rates—up to five times more intense—facilitating quicker turnover and responses to perturbations.^[21] Macrobenthos typically burrow or form visible structures on or in sediments, influencing larger-scale mixing, while meiobenthos' compact size and vagile (mobile) lifestyles restrict them to finer-scale activities without the extensive gallery-building seen in larger taxa. A key boundary challenge in meiobenthos classification involves the distinction between permanent meiofauna, which maintain their small size throughout their life cycles (e.g., nematodes and harpacticoid copepods), and temporary meiofauna, comprising juveniles or larval stages of macrobenthic species that only transiently fit the size criteria before growing larger.^[20] This overlap underscores that meiobenthos is not strictly a taxonomic group but a functional size class, with permanent forms exhibiting direct development and parthenogenesis for rapid reproduction, unlike the metamorphosis common in temporary members.^[22] Functionally, meiobenthos dominate pore water niches within sediments, where their movements facilitate microscale bioturbation, enhancing solute transport, nutrient cycling, and denitrification rates at the sediment-water interface—processes amplified by their high abundances and intimate contact with interstitial fluids.^[23] Unlike macrobenthos, which drive macroscale reworking, or microfauna limited to static microbial films, meiobenthos bridge these scales by grazing on bacteria and algae, thereby accelerating organic matter remineralization without relying on external bioturbators.^[24] Exclusion criteria for meiobenthos emphasize strictly benthic, vagile organisms; planktonic larvae or holoplanktonic forms are omitted despite similar sizes, as are predominantly sessile taxa that do not actively navigate sediments, ensuring the focus remains on interstitial dwellers integral to sediment dynamics. This delineation highlights meiobenthos' role as a discrete ecological entity, separate from epibenthic or pelagic components.^[25]

Historical Development

Origin of the Term

The term "meiobenthos" emerged from early 20th-century observations of small benthic organisms in marine sediments, particularly the interstitial fauna inhabiting the spaces between sand grains. In the 1930s, German zoologist Adolf Remane conducted pioneering studies on these communities in European coastal sands, such as those in the Kiel Bay, describing their distribution and ecological organization without yet proposing a unified nomenclature for the group. Remane's work highlighted the distinct biology of these minute, mobile animals in psammolittoral habitats, laying foundational insights into their adaptations to sediment interstices.^[26] The term itself was coined in 1942 by British marine biologist Mary F. Mare, commonly known as Molly Mare, in her seminal study of a benthic community in muddy sediments near Plymouth, England.^[27] Mare introduced "meiobenthos" to denote small metazoans—such as nematodes, copepods, and turbellarians—passing through a 1 mm mesh sieve but retained on finer meshes (typically 0.044 mm), distinguishing them from the larger macrobenthos and microbial components.^[27] Etymologically, it derives from the Greek meion (μείων, meaning "lesser" or "smaller") and benthos (βένθος, referring to the sea floor or depths), reflecting their intermediate size and habitat in benthic environments.^[28] This conceptualization arose from Mare's quantitative analysis of interstitial fauna in coastal sediments, emphasizing their role as a discrete ecological category separate from coarser macrofaunal groups.^[27] By the 1960s, "meiobenthos" became increasingly interchangeable with "meiofauna," a term also introduced by Mare in 1942, as researchers recognized the functional similarities across habitats.^[2] This terminological convergence was formalized through international collaborations, notably at the First International Conference on Meiofauna held in Tunis, Tunisia, from July 1-11, 1969, where participants standardized definitions and sampling methods for these small benthic invertebrates.^[29] The proceedings of this symposium underscored the term's applicability beyond marine contexts, promoting its use in comparative studies.^[29] The nomenclature's influence extended to limnology by the 1970s, where "meiobenthos" was adopted to describe analogous small-bodied communities in freshwater sediments, bridging marine and inland aquatic ecology. This expansion, detailed in comprehensive reviews like that of Remane and Schlieper, facilitated unified approaches to studying sediment-dwelling microfauna across diverse aquatic systems.^[30]

Key Milestones in Research

The study of meiobenthos gained momentum in the post-World War II era, building on the foundational work of Adolf Remane from the 1930s, who emphasized the high diversity and ecological significance of microscopic benthic organisms in marine sediments, laying the groundwork for recognizing meiofauna as a distinct functional group.^[26] Remane's investigations into interstitial fauna, including quantitative assessments of species richness in sandy habitats, highlighted the need for specialized approaches to studying these small metazoans beyond traditional macrofaunal surveys. During the 1960s and 1970s, the field solidified with international collaborations that established meiofauna as a dedicated research domain, exemplified by the First International Conference on Meiofauna held in Tunis in 1969, whose proceedings published in 1971 fostered global exchange on sampling and ecological roles.^[29] These efforts culminated in standardized size criteria, defining meiofauna as organisms retained on 40-63 μm sieves but passing through 0.5-1 mm meshes, as outlined in key manuals that promoted consistent methodologies across studies.^[2] Quantitative surveys during this period, such as those by A.D. McIntyre, advanced understanding of meiofaunal abundance and distribution in coastal sediments, integrating meiofauna into broader benthic ecology.^[31] In the 1980s, John S. Gray's syntheses on benthic ecology synthesized meiofaunal data into comprehensive frameworks, emphasizing their roles in sediment processes and community dynamics through influential texts like Ecology of Marine Sediments (1981).^[32] Milestone publications, including the second edition of Methods for the Study of Marine Benthos (1984) by N.A. Holme and A.D. McIntyre, provided standardized protocols for extraction and analysis, boosting comparative research worldwide.^[33] The 1980s and 1990s saw advances in ultrastructural and molecular techniques, with scanning electron microscopy (SEM) revealing fine morphological details of nematodes, such as cuticular structures and sensory organs, enhancing taxonomic precision.^[34] Early molecular studies, including DNA sequencing of ribosomal genes, uncovered cryptic species diversity within nematode genera, challenging prior morphological classifications and indicating higher biodiversity than previously estimated.^[35] From the 2000s onward, metagenomic approaches integrated environmental DNA sequencing to assess meiobenthic community composition without culturing, revealing previously undetected microbial interactions and diversity patterns in sediments.^[36] Concurrently, studies on polar meiobenthos in the 2010s documented responses to climate warming, such as shifts in nematode and copepod abundances linked to sea ice reduction and temperature rises in Antarctic and Arctic regions.^[37] In the 2020s, continued advances in metabarcoding and eDNA techniques have further elucidated meiobenthos biodiversity and responses to environmental changes, including ocean acidification and pollution, as of 2025.^[38] Key texts like Introduction to the Study of Meiofauna (1988) by R.P. Higgins and H. Thiel, and the second edition of Meiobenthology (2009) by O. Giere, consolidated these developments, offering updated techniques and ecological syntheses that continue to guide the field.^[39]^[40]

Taxonomic Composition

Major Taxonomic Groups

Meiobenthos communities are dominated by metazoans, with the majority of meiofauna biomass contributed by organisms from approximately 20 metazoan phyla, primarily those fitting the operational size range of 45-1000 μm. Key groups include Nematoda, which often comprise the majority of individuals; Arthropoda, represented mainly by harpacticoid copepods and ostracods; Annelida, including small polychaetes and oligochaetes; Platyhelminthes (turbellarians); Gastrotricha; Rotifera; Tardigrada; and Kinorhyncha.^[4]^[40] Protozoan components, such as Foraminifera, ciliates, and other sarcodines (e.g., testate amoebae), are frequently incorporated into meiobenthos assessments due to their comparable dimensions and benthic habitat occupancy, though they are unicellular and ecologically distinct from metazoans.^[4]^[41] Less prevalent taxa, such as Cnidaria (particularly hydrozoans), Nemertea, and Priapulida, constitute less than 5% of overall meiobenthos abundance and biomass across diverse sediments.^[4] Taxonomic classification of meiobenthos faces significant hurdles due to pronounced endemism, cryptic speciation, and extensive under-sampling in remote habitats, leading to conservative global species estimates surpassing 100,000.^[42]^[43] Standard overviews of these groupings draw from foundational reviews, including Higgins and Thiel (1988) for methodological and systematic foundations, and Danovaro et al. (2013) for updated compositional patterns in coastal settings.^[44]^[45]

Dominant Taxa and Diversity Patterns

Nematodes are the dominant taxon within meiobenthic communities, typically comprising 50-90% of total meiofaunal abundance and often exceeding 80% in many marine sediments.^[46] These free-living species exhibit diverse feeding strategies, including bacterivory and omnivory, which enable them to exploit a wide range of microbial and detrital resources in interstitial spaces.^[46] Nematode species richness is notably high, with hundreds of species documented per site in deep-sea environments, reflecting their adaptive radiation across varied sediment conditions.^[47] Harpacticoid copepods represent the second most abundant group, accounting for 10-30% of meiobenthic individuals in coastal and shelf sediments.^[48] These arthropods include both epibenthic forms that crawl over sediment surfaces and interstitial species that navigate pore spaces, contributing significantly to secondary production through their grazing on microalgae such as diatoms.^[49] Their role in carbon transfer underscores their importance in benthic food webs, where they convert primary production into biomass available to higher trophic levels.^[50] Meiobenthic diversity patterns show elevated levels in shallow, organic-rich sediments, where enhanced nutrient availability supports greater taxonomic richness and abundance compared to deeper or nutrient-poor areas.^[51] Latitudinal gradients reveal peaks in the tropics, with species diversity declining toward polar regions, consistent with broader marine biodiversity trends driven by temperature and productivity variations.^[46] Studies employing DNA barcoding have uncovered cryptic speciation, indicating that morphological assessments underestimate true diversity, particularly among nematodes and copepods.^[52] Despite their small size, nematodes often contribute more than 50% of total meiofaunal biomass, highlighting their disproportionate ecological weight in sediment ecosystems.^[46] Diversity exhibits variability across environmental gradients, with reduced species richness in hypoxic or polluted sediments; harpacticoid copepods prove more sensitive than nematodes, often showing sharper declines in abundance under low-oxygen conditions.^[53]^[54]

Habitats and Distribution

Sediment Environments

Meiobenthos primarily inhabit the interstitial pore spaces within aquatic sediments, navigating networks formed by grains typically ranging from 63 to 500 μm in diameter, which align with their body sizes of 32 μm to about 1 mm. This lifestyle demands specific adaptations, including slender, flexible bodies for maneuvering in confined areas under elevated hydrostatic pressure and tolerance to low oxygen levels through mechanisms like anaerobic metabolism or vertical migration. Such conditions are common in subsurface sediments, where dissolved oxygen diminishes rapidly, yet many meiobenthic taxa persist by exploiting micro-oxic niches or sulfidic environments.^[1]^[55]^[56] These organisms predominate in fine sands, silts, and muds, which offer abundant, interconnected interstices conducive to high population densities, while avoiding coarse gravels and pebbles due to sparse and unstable pore spaces that limit habitat suitability. Sensitivity to grain size is pronounced, with optimal conditions in the phi scale range of 2–4 (corresponding to 0.25–0.063 mm, or fine sand to coarse silt), where porosity balances permeability and retention of organic particles. Empirical models link sediment porosity—often 20–40% in these grain sizes—to meiobenthos abundance, showing positive correlations as higher porosity facilitates oxygen diffusion and food access.^[57]^[58]^[59] Microscale chemical conditions critically influence meiobenthos viability, including oxygen penetration depths of 1–5 mm into the sediment, beyond which anoxia prevails and restricts most taxa. Organic content, typically 1–3% in productive habitats, provides essential bacterial-derived food resources, while salinity gradients—ranging from freshwater to full marine—shape community structure by affecting osmotic regulation and microbial activity. Vertical stratification reflects these gradients: oxygenated species cluster in surface layers (0–1 cm), whereas hypoxia-tolerant nematodes extend into deeper burrows up to 10 cm, accessing alternative electron acceptors. Dominant taxa like nematodes particularly thrive in these fine sediments, underscoring their adaptability.^[56]^[59]^[60]^[61]

Global Occurrence and Adaptations

Meiobenthos exhibit a pronounced dominance in marine environments, where they are ubiquitous across a wide range of depths and habitats, from intertidal zones to abyssal plains exceeding 4000 meters. Highest densities are typically observed in coastal and continental shelf areas, where organic matter availability supports abundant populations, often reaching thousands of individuals per 10 square centimeters in sediments.^[62] In contrast, their presence in freshwater and brackish systems, such as lakes, rivers, and estuaries, is less diverse but still significant, with communities adapted to variable conditions like fluctuating salinity through euryhaline capabilities, particularly in nematodes that can tolerate salinities from near-zero to over 50 practical salinity units.^[38] For instance, nematodes and harpacticoid copepods dominate these transitional zones, facilitating their persistence in dynamic estuarine gradients.^[63] Meiobenthos also inhabit extreme environments, including polar ice sediments, hydrothermal vents, and hypersaline lagoons, demonstrating remarkable tolerances to harsh conditions such as temperatures from -2°C to 40°C and prolonged anoxia. In polar regions, nematodes like Monhystera species thrive in sub-zero sediments by employing dormancy mechanisms to endure low temperatures and ice cover, while in hydrothermal vents, taxa such as Oncholaimus nematodes and dirivultid copepods withstand temperatures up to 40°C, low oxygen, and high sulfide levels through sulfur detoxification and enhanced oxygen-binding proteins like haemoglobin.^[55] Hypersaline lagoons host specialized forms, including loriciferans (Spinoloricus cinziae) in anoxic brines exceeding 200 practical salinity units, relying on symbiotic sulfur-oxidizing bacteria and hydrogenosome-based metabolism for survival under extreme osmotic and hypoxic stress.^[55] These adaptations enable meiobenthos to colonize niches otherwise inhospitable to larger biota.^[64] Biogeographic patterns of meiobenthos reveal a mix of cosmopolitan and regionally endemic species, with nematodes showing widespread distribution due to high dispersal potential via dormant stages, yet many genera exhibit habitat-specific endemism. While some taxa like Sabatieria and Halalaimus nematodes appear cosmopolitan across oceans, genetic analyses often uncover cryptic species complexes masking true endemism, particularly in isolated systems like Lake Baikal, where 19 harpacticoid species are endemic.^[65] Deep-sea diversity is notably lower, attributed to food limitation from reduced organic flux, with expected genus richness dropping from around 37 at 500 meters to 21 beyond 4000 meters, alongside increased singleton taxa indicating potential undiscovered endemics.^[65]^[63] Key physiological adaptations underpin the global success of meiobenthos, including cuticular flexibility in nematodes that allows undulatory locomotion through tight sediment pores, chemosensory organs such as amphids for detecting resources and chemical cues in opaque environments, and rapid osmoregulation via active ion transport to maintain internal balance amid salinity shifts.^[66] These traits, combined with dormancy for anoxic survival and symbiotic associations for nutrient acquisition, enable efficient navigation, resource exploitation, and resilience across diverse ecosystems.^[55]^[67]

Ecological Roles

Trophic Interactions

Meiobenthos exhibit diverse feeding guilds that position them as key consumers in benthic food webs, primarily targeting particles in the size range of 1-50 μm, such as bacteria, microalgae, and detritus. Bacterivory is the dominant mode among nematodes, which comprise the majority of meiobenthic taxa and selectively ingest bacteria via deposit-feeding mechanisms classified as Type 1A based on buccal cavity morphology. Herbivory occurs through grazing on microalgae and diatoms by epistrate feeders (Type 2A), while carnivory is observed in omnivorous or predatory nematodes (Type 2B) that consume smaller meiofauna like protozoans or juvenile nematodes. Dominant taxa such as harpacticoid copepods also function as grazers on microalgae, contributing to these interactions.^[68]^[69] As prey, meiobenthos serve as a primary food source for higher trophic levels, including juvenile fish species like carps and gudgeons, as well as macroinvertebrates such as chironomid larvae, plecopterans, and crustaceans. Gut content analyses reveal high ingestion rates, with juvenile fish consuming up to 234,000 nematodes per day and macroinvertebrates like flatworms ingesting hundreds of nematodes in hours, underscoring their nutritional importance in estuarine and coastal systems. Trophic transfer efficiency to macrobenthos is estimated at approximately 10-20%, reflecting partial energy passage through predation despite high meiofaunal turnover.^[70] Symbiotic and parasitic interactions further integrate meiobenthos into benthic networks, with many nematode genera acting as endoparasites within larger hosts like macroinvertebrates or fish, exploiting host tissues for nutrition. Mutualistic associations with bacteria aid digestion, particularly in nematodes where gut symbionts enhance breakdown of refractory organic matter.^[1]^[71] In trophic structure models, meiobenthos act as an intermediate link between microbial communities and macrofauna, channeling energy from primary producers like microphytobenthos (contributing 60-81% of consumption) and bacteria to higher levels via short, efficient pathways. Their high turnover rates result in annual biomass production 10-100 times that of macrobenthos, driven by elevated production-to-biomass ratios (0.03-0.28 d⁻¹ versus <0.01 d⁻¹ for macrofauna), as quantified in linear inverse food web models. Isotopic studies using δ¹³C and δ¹⁵N signatures confirm their detritivorous roles in coastal systems, with values indicating primary consumer positions (δ¹⁵N: 4-8‰) and reliance on mixed autochthonous-allochthonous carbon sources (δ¹³C: -38 to -30‰), particularly in detritus-dominated habitats.^[72]^[73]

Ecosystem Functions and Indicators

Meiobenthos mediate the remineralization of carbon and nitrogen in benthic ecosystems primarily through grazing on organic detritus and microbial biofilms, as well as via nutrient excretion that recycles bioavailable forms back into the sediment-water interface.^[25] Their bioturbatory activities enhance these processes by increasing solute transport rates by factors of 1.5 to 3.1, thereby stimulating microbial decomposition of organic matter.^[74] In anoxic sediments, meiobenthos significantly boost denitrification by promoting oxygen and nitrate diffusion to denitrifying bacteria, with enhancements reaching up to 50% of total benthic nitrogen flux in high-density assemblages.^[23] Through small-scale reworking of sediment particles, meiobenthos facilitate bioturbation that extends oxygen penetration depths by 59–85% in hypoxic environments, counteracting sulfide accumulation and stabilizing sediment structure via mucus-lined burrows.^[75] This increased oxygenation supports elevated microbial activity, including aerobic respiration and sulfate reduction, which collectively improve organic matter breakdown and prevent anoxic collapse in coastal sediments.^[25] Additionally, meiobenthos serve as an important prey base for larger benthic and demersal fish, thereby integrating nutrient flows into higher trophic levels. Meiobenthos fulfill a keystone role in sustaining microbial biodiversity by exerting top-down control through selective grazing, which maintains bacterial and diatom populations in active growth phases and prevents dominance by opportunistic species.^[25] This grazing pressure, combined with enzymatic contributions from meiofaunal mucus, fosters diverse microbial assemblages essential for ecosystem resilience and primary production in sediments. As environmental sentinels, meiobenthos exhibit sensitivity to pollutants like heavy metals, which disproportionately affect copepods over nematodes, leading to elevated nematode:copepod ratios that signal contamination levels in biomonitoring efforts.^[76] Such ratios are widely applied in marine health assessments to detect anthropogenic stress. Regarding climate impacts, ocean acidification alters meiobenthic community structure by favoring tolerant predatory nematodes while reducing overall functional diversity, with models projecting 20–50% diversity losses in continental shelf habitats by 2100 under high-emission scenarios.^[77]

Collection and Analysis Methods

Sampling Techniques

Sampling techniques for meiobenthos are designed to collect small, sediment-dwelling organisms (typically 20–500 μm in size) while minimizing disturbance to the sediment structure and preserving vertical distributions. These methods vary by habitat depth and type, with core samplers preferred for quantitative assessments in soft sediments to maintain intact layers.^[78]^[79] In shallow waters, such as intertidal or subtidal zones up to a few meters deep, hand corers made from 10 cm diameter PVC tubes are commonly used to extract undisturbed sediment cores. These manual devices allow precise penetration to depths of 10–20 cm, capturing the upper sediment layers where most meiobenthos reside, and are ideal for soft muds or sands in accessible areas.^[78] For deeper marine environments, multicorers equipped with multiple 10 cm inner diameter tubes (e.g., 4–12 tubes per deployment) or box corers (0.25–0.5 m² surface area) are deployed from research vessels to sample bathyal and abyssal depths, preserving overlying water and sediment interfaces critical for meiofaunal studies. Multicorers provide higher quantitative accuracy for abundance estimates compared to box corers, which are better suited for qualitative diversity assessments due to their larger sampled volume.^[79]^[80] Grab samplers, such as the Van Veen (0.1 m²) or Ponar grabs, are employed for broader coverage in soft sediments across continental shelves and slopes, though they disrupt vertical profiles and are less suitable for precise depth-specific sampling. These clamshell devices penetrate 10–20 cm into the seabed, after which subsamples (e.g., via mini-corers) are taken onboard to target meiobenthos, with Van Veen grabs preferred for their reliability in fine-grained substrates.^[81] For inaccessible or visually guided collection, diver-operated methods using SCUBA in shallow coastal zones (up to 30 m) involve hand-held corers or suction samplers to target specific microhabitats like seagrass beds. In deep-sea settings, remotely operated vehicles (ROVs) facilitate targeted sampling with push corers or suction devices, allowing undisturbed collection from chemosynthetic habitats or hard substrates where traditional gear fails. Quantitative pontoon samplers, deployed from floating platforms, are used in freshwater lakes to collect replicate cores from profundal zones.^[81]^[82] Standard protocols emphasize 3–5 replicates per site to account for spatial patchiness, with each sample covering 0.01–0.1 m² to adequately represent density variability (typically 10³–10⁶ individuals m⁻²).^[79] In the field, samples are immediately fixed in 4–10% buffered formalin or 70–95% ethanol to halt degradation and maintain specimen integrity for subsequent processing, with formalin preferred for morphological studies and ethanol for molecular analyses.^[83]^[84]

Extraction and Identification Procedures

Extraction of meiobenthos from sediment samples typically begins with mechanical sieving to separate organisms based on size. Sediment is sequentially passed through 500 μm and 45 μm mesh sieves using filtered seawater to retain meiofaunal organisms while discarding larger macrofauna and finer particles.^[85]^[79] During this process, the sample is agitated—often by gentle stirring or swirling—to dislodge organisms adhered to sediment grains, followed by decantation where the supernatant containing suspended fauna is collected and sieved.^[2] This method achieves extraction efficiencies of 80-90% for most taxa in sandy or muddy substrates, though it may underestimate abundances in cohesive silty muds due to incomplete release.^[84] For enhanced recovery, particularly in fine-grained sediments, chemical extraction techniques exploit density differences between organisms and particles. Ludox-TM, a colloidal silica solution with a specific gravity of approximately 1.13, is commonly used for flotation: sediment is mixed with diluted Ludox-TM, allowed to settle briefly, and the floating organic fraction containing meiobenthos is decanted or centrifuged at 1800 rpm for 10 minutes to concentrate specimens.^[86]^[87] To facilitate live sorting, a 6% MgCl₂ solution in seawater is added as an anesthetic, relaxing motile organisms without immediate fixation and allowing separation under low magnification.^[85] These procedures, often combined with sieving, improve yields for nematodes and copepods, key meiobenthic groups, by minimizing loss in dense sediments.^[88] Once extracted, specimens are stained and sorted for analysis. Rose Bengal dye (1 g/L in preservative) is applied to preserved samples to enhance contrast of soft-bodied organisms against sediment residue, enabling visualization under a stereomicroscope at 25-50× magnification.^[2]^[84] Live versus dead discrimination occurs through observation of movement in anesthetized or unstained aliquots, with viable specimens pipetted into separate vials using fine tools like silicone-coated Pasteur pipettes.^[85] Lignin or other vital stains may supplement Rose Bengal for specific taxa, though care is taken to avoid over-staining that could obscure morphological details.^[2] Identification relies on morphological examination, particularly for dominant groups like nematodes, which are often identified to genus using specialized keys that emphasize features such as amphids, stoma structure, and tail shape.^[84] Specimens are relaxed with heat (60-70°C) or MgCl₂, fixed in 4-5% buffered formalin for 24 hours, and mounted on glass slides in glycerin or lactophenol for high-resolution light microscopy at 400-1000×.^[85]^[2] For other taxa like harpacticoid copepods, phase-contrast microscopy aids in viewing appendages and segmentation. Emerging molecular approaches, such as DNA metabarcoding, complement traditional methods by extracting DNA from bulk sediment (5-10 g) or sorted meiofauna using kits like QIAGEN PowerSoil, followed by PCR amplification of the 18S rRNA gene (e.g., V1-V2 region with primers SSU_F04/SSU_R22) and Illumina MiSeq sequencing for operational taxonomic unit assignment.^[89] This technique accelerates biodiversity assessment but requires bioinformatics pipelines to resolve cryptic species, often validating results against morphological identifications.^[89] Quantification standardizes abundances as individuals per 10 cm³ of processed sediment volume, derived from core dimensions and subsample fractions to ensure representativeness.^[84] Biomass is measured via wet weight (blotted and weighed to 0.01 mg accuracy) for fresh or preserved samples, or through CHN elemental analysis after drying and grinding pooled specimens, providing carbon-based estimates that account for taxonomic composition.^[85] These metrics support comparisons across studies, with extraction efficiency verified by re-examination of residues.^[2]

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The meiofauna are defined as those benthic animals that pass through a 500-μm sieve but are retained on a 40-μm sieve (Fenchel 1978, Higgins and Thiel 1988).Missing: scholarly | Show results with:scholarly
[15]
Should we redefine meiofaunal organisms? The impact of mesh size ...
They are defined by mesh size and specifically by their retention on a sieve with a 44-µm mesh size. In this study, we examined the accuracy of this ...
[16]
Abundance and Biomass of Metazoan Meiobenthos in The Deep Sea
Abundance and biomass in the deep Atlantic, Pacific and Indian Oceans generally range between 100 and 1000 x 103 individuals m-2 and 10 to 400 mg C m-2, ...Missing: per m2
[17]
[PDF] On the importance of marine meiofauna for benthos communities
Thiel (1966) counted in deep sea sediment from the Indian Ocean (3000-5000 m) be tween 20000 and 110000 meiofauna individuals per m2. If one assumes a wet ...
[18]
Biomass and production of freshwater meiofauna - ResearchGate
Aug 6, 2025 · As nematodes have fast turnover rates (generation times of a few days or weeks), it is often difficult to have a sampling frequency adapted ...
[19]
Benthic community patterns and the composition of feeding-types ...
Aug 10, 2025 · Organisms reproducing by cyclical parthenogenesis combine the benefits of both sexual and asexual reproduction within the same life cycle.
[20]
Meiofauna of Sandy Beaches - Coastal Wiki
May 7, 2025 · Nowadays the term meiofauna is used interchangeably with meiobenthos. Meiofauna is mainly found in and on soft sediments, but also on underwater ...
[21]
[PDF] Chapitre VIII Macrobenthos and meiobenthos
It seems justified to assume that metabolism in meiobenthos is 5 times more intense than in macrobenthos. Besides, biomass of meiobenthos is generally ...
[22]
(PDF) Meiobenthos and macrobenthos are discrete entities and not ...
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Meiofauna increases bacterial denitrification in marine sediments
Oct 16, 2014 · Denitrification is double that in sediments with low meiofauna, suggesting that meiofauna bioturbation has a stimulating effect on nitrifying and denitrifying ...<|control11|><|separator|>
[24]
Meiobenthos in marine coastal sediments - GeoScienceWorld
Sep 16, 2013 · Meiobenthic organisms are inter- mediate in size between macro- and microfauna, and their size limits are defined based on standar- dized mesh ...
[25]
Meiofauna matters: The roles of meiofauna in benthic ecosystems
Aug 7, 2025 · macrofauna more often exhibit a lifestyle whereby they rely on sediment displacement or burrowing. The highly conservative bimodal body size ...
[26]
(PDF) Bryozoans of the Baltic Sea - ResearchGate
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[PDF] REFERENCE TO THE MICRO-ORGANISMS By :MollyF. Mare
It has been necessary to propose the following new terminology: the. Macrobenthos, which is here equivalent to the macrofauna, the Meiobenthos, under which term ...
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https://www.oed.com/dictionary/meiobenthos_n?tl=true
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A study of a marine benthic community with special reference to the ...
May 11, 2009 · A quantitative ecological study of the microbenthos has been started and the habitat is first described, stress being laid on points of ...
[30]
[PDF] Proceedings of the First International Conference on Meiofauna
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Proceedings of the First International Conference on Meiofauna
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Biology of Brackish Water - Adolf Remane, Carl Schlieper
Adolf Remane, Carl Schlieper. Edition, 2, illustrated. Publisher, E. Schweizerbartʹsche Verlagsbuchhandlung, 1971. Original from, the University of Virginia.Missing: Meiobenthology pdf
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(PDF) Adolf Remane (1898-1976) and his views on systematics ...
Aug 6, 2025 · PDF | Adolf Remane was primarily a morphologist and systematist. In 1952, he published an influential book on the foundations of systematics ...
[34]
[PDF] Progress in Oceanography
The term meiofauna was introduced and defined by Mare (1942). It derived from the Greek word 'meio' meaning 'the smaller'. Quantitative studies on sediment-.Missing: origin Remane
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N. A. Holme & A. D. Mcintyre, editors. Methods for the Study of ...
N. A. Holme & A. D. Mcintyre, editors. Methods for the Study of Marine Benthos. 2nd edition, xii, 387 pp. Blackwell, 1984. Price £25.00.<|separator|>
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N. A. Holme and A. D. McIntyre (Editors): Methods for the Study of ...
N. A. Holme and A. D. McIntyre (Editors): Methods for the Study of Marine Benthos. Second Edition = IBP Handbook 16.–399 pp. Oxford-London-Boston: Blackwell ...
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[PDF] biodiversity, distribution patterns and trophic position
SCANNING ELECTRON MICROSCOPY. Any ectosymbiotic relation in the nematodes was checked with scanning electron microscopy. (SEM). Nematode specimens were fixed ...
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[PDF] A critique of current methods in nematode taxonomy
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Sediment DNA metabarcoding and morphology provide ...
In this study we analyse the effects of glacially mediated disturbance on the biodiversity of benthic macrofauna and meiobenthos (meiofauna and Foraminifera) ...
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Spatial and temporal dynamics of Antarctic shallow soft-bottom ...
Jul 1, 2019 · This is particularly true of the Polar Regions, which are currently undergoing rapid climate change, the impacts of which are poorly understood.
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Meiobenthology: The Microscopic Motile Fauna of Aquatic Sediments
Free delivery 14-day returnsMeiobenthology is the science of the tiny animals that live in huge numbers in all aquatic sediments. This fully revised and enlarged second edition ...
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Metazoan meiobenthos along continental margins: a review
This review compiles abundance and biomass data from a total of 38 papers, describing metazoan meiofauna assemblages along the continental margins.
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Guide to the Identification of Marine Meiofauna - Allen Press
Nov 18, 2020 · There are earlier books on meiofauna (Hulings & Gray 1971, Higgins & Thiel 1988, Giere 2009) that provide good overviews but leave gaps in ...
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WoRMS source details - WoRMS - World Register of Marine Species
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(PDF) Reevaluation of species diversity patterns of free-living ...
Aug 5, 2025 · Nematode species diversity was analysed for 55 data points ranging from local studies to large-scale faunal surveys.
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Meiofaunal communities in a tropical seagrass bed and adjacent ...
Nematoda was the most abundant taxon (72.8%) in the seagrass quadrats, followed by Copepoda (13.7%) and Polychaeta (6.1%). However, Copepoda accounted for 45.2 ...
[51]
Productivity of bacteria and microalgae and the effect of grazing by ...
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[52]
Grazing on diatoms by harpacticoid copepods: species-specific ...
Harpacticoid copepods are an important part of the meiofauna, consuming pri- mary production in planktonic, benthic and epiphytic communities (Hicks & Coull ...
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Organic biomarkers and Meiofauna diversity reflect distinct carbon ...
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Barcoding against a paradox? Combined molecular species ...
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Organic enrichment negatively impacts meiobenthic copepods on ...
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Effects of Hypoxia and Anoxia on Meiofauna: A Review with New ...
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Characteristics of meiofauna in extreme marine ecosystems: a review
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Meiofauna improve oxygenation and accelerate sulfide removal in ...
Unlike macrofauna, meiofauna are abundant in hypoxic sediments. We studied to what extent meiofauna affect oxygen availability, sulfide removal and microbial ...
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Meiobenthology - The Microscopic Fauna in Aquatic Sediments
Studies on meiobenthos, the motile microscopic fauna of aquatic sediments, are gaining in importance, revealing trophic cycles and assessing impacts of anthro-.
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(PDF) Chapter 14 - Meiofauna. - ResearchGate
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Vertical gradients of meiofauna and bacteria in subtidal sandy ...
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Abundance and size distribution of sublittoral meiobenthos along ...
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[63]
[PDF] Vertical distribution of meiobenthos in the sediment profile in bathyal ...
Organisms were obviously concentrated in the surface 0-1 cm layer of the sediment, and decreased exponentially in number with increase of depth in the sediment.
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Meiofauna: Biodiversity, Ecology, and Role in Ecosystems - MDPI
Sep 1, 2023 · Meiobenthic communities are characterized by a smaller scale variability of structure than macrozoobenthos communities, and seasonal changes in ...
[65]
Fundamental questions in meiofauna research highlight how small ...
Mar 17, 2025 · This paper identifies the top-50 priority questions for meiofaunal research, highlighting their critical roles in biogeochemical cycles and biodiversity.
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Meiofauna—Adapted to Life at the Limits - Archimer
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Cosmopolitism, rareness and endemism in deep-sea marine ...
Amongst meiofaunal organisms, nematodes are expected to have a high species richness, with an estimated ca 50,000 to >100,000 species (based on expert ...
[68]
Nematodes ultrastructure: complex systems and processes - PMC
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(PDF) Nematodes in aquatic environments: adaptations and survival ...
Apr 11, 2015 · They represent great degree of morphological adaptations including those associated with cuticle, sensilla, pseudocoelomic inclusions, stoma, ...
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[PDF] Do meiofauna consume primary production?
What do meiofauna feed on? As a group, meiofauna consume a wide variety of food sources, including detritus, bacteria, diatoms and other small photoautotrophs, ...
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[PDF] 6 Feeding Ecology of Free-living Benthic Nematodes
This chapter aims to give an overview of feeding habits and food sources of aquatic nematodes, and touches upon issues of food recognition, feeding selectivity,.
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Are meiofauna a standard meal for macroinvertebrates and juvenile ...
Feb 14, 2020 · Here we present a synthesis of recent evidence showing that meiofauna are significant as prey, not only for many benthic macroinvertebrates (chironomids, ...
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Master of Puppets: How Microbiota Drive the Nematoda Ecology ...
Aug 19, 2025 · The nematode microbiota can influence life-history traits, thereby shaping the species' ecology and evolutionary trajectories.
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Quantitative food web modeling unravels the importance of the ...
Meiofauna production was higher than those of macrofauna in mudflat MO, seagrass SR and sandflat SR, and relatively similar in mudflat SR and seagrass MO.
[75]
Trophic positioning of meiofauna revealed by stable isotopes and ...
Aug 6, 2016 · Meiofauna are capable of extremely high biomass turnover times with up to 1.08 d (i.e., Stead et al. 2005), thus, to ensure better stable ...Methods · Results · Meiofauna Isotope Analysis<|separator|>
[76]
Marine meiofauna, carbon and nitrogen mineralization in sandy and ...
Feb 21, 2025 · ABSTRACT: Organic carbon mineralization was studied in a shallow-water (4 m), sandy sediment and. 2 comparatively deep-water (150 and 300 m) ...
[77]
Meiofauna improve oxygenation and accelerate sulfide removal in ...
Temporary hypoxic-anoxic sediments at these depths have meiofauna abundances of 24–3452 ind. ... Meiofauna increases bacterial denitrification in marine sediments.
[78]
Nematode/copepod ratio and nematode and copepod abundances ...
Jul 20, 2022 · Based on different pollution tolerance, a nematode/copepod ratio was proposed as an easy tool for monitoring the effect of anthropogenic ...
[79]
Meiobenthos and ocean acidification: Effects on meiobenthic ...
Overall, the present study revealed a relatively high tolerance of meiobenthos and nematodes to lower pH conditions. However, an in-depth analysis of nematode ...
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Sampling tools for the marine environment - Coastal Wiki
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Comparison of sampling methods for deep‐sea infauna - ASLO - Wiley
Nov 21, 2016 · Multicorers are recommended for quantitative assessments, but box corers are useful for qualitative studies that require capturing more ...
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[PDF] A multiple corer for taking virtually undisturbed samples from shelf ...
If the initial sampling is in shallow water the corer should first be suspended in the sea for a few minutes to allow the empty hydraulic damper cylinder to ...Missing: multicorers | Show results with:multicorers
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[PDF] Soft bottom macrofauna: Collection and treatment of samples
For descriptive surveys, sieves used for extraction of the macro- fauna from sediments should have a mesh size of 1.0 mm. The use of an additional finer sieve ...Missing: criteria operational scholarly
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[PDF] THE MEIOBENTHOS OF THE MOLLOY DEEP (5 600 M), FRAM ...
Apr 22, 2021 · In 1999, the ROV allowed targeted sampling of areas of the seabed that appeared visually to be perfectly undisturbed,. i.e. also without any ...
[85]
Cost/benefit and the effect of sample preservation procedures on ...
Oct 27, 2017 · The commonly used sample preservation procedures are: (1) fixation in 4–10% formalin and subsequent preservation in 70% ethanol; or (2) simply ...Missing: meiobenthos replication
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(PDF) Methods for quantitative estimation of meiobenthos
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[PDF] METHODS FOR THE STUDY OF MARINE BENTHOS
... Methods for the study of marine benthos / edited by Anastasios Eleftheriou, Hellenic Centre for Marine. Research, Crete, Greece and Department of Biology ...
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An improved protocol for separating meiofauna from sediments ...
Apr 26, 2025 · The method uses colloidal silica Ludox®. (DuPont) HS 40 as a high density solution to extract meiofaunal organisms from sediment. Previous.
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Suggested criteria for describing deep-sea benthic communities
Meiofauna was extracted by flotation and centrifugation (1800 rpm, 10 min) using a colloidal silica solution (LudoxTM, Aldrich Chemical Company) with a specific ...
[90]
An improved protocol for separating meiofauna from sediments ...
Aug 5, 2025 · The method uses colloidal silica Ludox((R)) (DuPont) HS 40 as a high density solution to extract meiofaunal organisms from sediment. Previous ...
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[PDF] DNA Metabarcoding Methods for the Study of Marine Benthic ...
Sep 30, 2021 · We identified several key aspects that should receive attention in future studies on marine meiobenthos: (1) comparison of sampling methods, ...