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Nereis

Nereis is a genus of polychaete annelids in the family Nereididae, established by Linnaeus in 1758, and serves as the type genus for the family, with approximately 226 species currently attributed to it (as of 2023), though many may belong to other genera due to ongoing taxonomic revisions, primarily distributed in marine environments worldwide.^[1] These worms are characterized by elongated, segmented bodies comprising numerous segments (typically 100–200), paired parapodia bearing chaetae for locomotion and respiration, an eversible muscular pharynx armed with toothed jaws for predation, and paragnaths on the proboscis for sensory and feeding functions.^[1]^[2] They inhabit diverse benthic habitats from intertidal zones to abyssal depths, often in soft sediments like sand or mud, where they burrow, crawl, or swim, and some species tolerate estuarine or even freshwater conditions with fluctuating salinities.^[1]^[2] Ecologically significant as predators or omnivores feeding on small invertebrates, algae, or detritus, Nereis species are key components of food webs, serving as prey for fish and birds, and are studied in aquaculture and as bioindicators due to their sensitivity to environmental changes.^[2] A defining feature is their life history involving epitoky, where non-reproductive atokous forms transform into sexual epitokous stages with enlarged eyes and modified swimming appendages for synchronized spawning, often triggered by lunar or tidal cycles.^[1]^[2]

Taxonomy and Etymology

Classification

Nereis is classified within the kingdom Animalia, phylum Annelida, class Polychaeta, order Phyllodocida, family Nereididae, and genus Nereis Linnaeus, 1758.^[3] This placement reflects its status as a polychaete worm with segmented body structure and bristle-like setae, typical of errantian annelids. The genus was originally described by Carl Linnaeus in his Systema Naturae (10th edition), encompassing marine and estuarine species known for their active locomotion.^[3] The type species for Nereis is Nereis pelagica Linnaeus, 1758, designated by subsequent designation by Fauchald (1977).^[3] Historically, the genus has accumulated several synonyms, including Heteronereis Örsted, 1843 (a junior subjective synonym referring to epitokous stages), Lycoris Lamarck, 1818, Johnstonia Quatrefages, 1850 (suppressed by ICZN Opinion 1807), Naumachius Kinberg, 1865, and Thoosa Kinberg, 1865 (homonym and subjective synonym).^[3] These synonyms arose from early descriptions of heteronereid (swarming) forms, which were once thought to represent distinct genera but are now recognized as modified adults within Nereididae.^[3] Subgenera within Nereis have undergone significant revisions, with many elevated to separate genera based on morphological and molecular evidence. Notable historical subgenera include Nereis (Neanthes) Kinberg, 1865, Nereis (Ceratonereis) Kinberg, 1865, Nereis (Hediste) Malmgren, 1867, and Nereis (Platynereis) Kinberg, 1865, distinguished by variations in paragnath patterns, setal morphology, and reproductive modifications.^[3] Molecular phylogenetic studies using mitochondrial and nuclear DNA have revealed paraphyly in traditional groupings like Nereidinae, prompting reclassifications; for instance, species formerly in Nereis (Hediste) are now placed in Hediste (e.g., Hediste diversicolor), and those in Nereis (Platynereis) in Platynereis (e.g., Platynereis dumerilii), supported by analyses showing distinct clades within Nereididae.^[3]^[4] These revisions emphasize the role of genetic data in resolving cryptic diversity and evolutionary relationships. As of 2025, the genus comprises approximately 220 accepted species, with recent additions such as Nereis quadrata (Bonyadi-Naeini et al., 2025) and ongoing refinements based on molecular studies (Read & Fauchald, 2023).^[3]^[5] Key diagnostic features for identifying the genus Nereis include biramous parapodia on most segments for locomotion and respiration, heterogomph falcigerous setae (with composite falcigers in both rami and simple hooded hooks posteriorly), and a eversible proboscis equipped with four longitudinal rows of paragnaths on the maxillary ring and a pair of robust, fang-like jaws with serrated edges.^[3]^[6] These traits distinguish Nereis from related genera like Neanthes (lacking falcigers in certain positions) and Platynereis (with more uniform setal arrangements), aiding in taxonomic delineation despite ongoing refinements from molecular studies.^[3]

Etymology and Naming History

The genus name Nereis is derived from the Greek term "Nēreïs" (Νηρηΐς), referring to the sea nymphs of Greek mythology known as the Nereids, which aptly reflects the predominantly marine habitat of the organisms within this genus.^[7] The genus was first formally established by Carl Linnaeus in his seminal work Systema Naturae (10th edition), where he included five species under Nereis, with Nereis pelagica later designated as the type species by subsequent taxonomic authorities.^[8] This initial description marked Nereis as a foundational taxon in annelid classification, encompassing a broad array of polychaete worms characterized by their errant lifestyle and bristle-bearing appendages. In its early history, the genus Nereis was broadly circumscribed, incorporating numerous polychaete species that shared superficial morphological traits such as parapodia and setae, leading to significant taxonomic lumping.^[7] Key early delineations came from naturalists like Henri Marie Ducrotay de Blainville, who in 1818 established the family Nereididae (to which Nereis belongs) and contributed species descriptions that helped refine genus boundaries in the 1820s.^[9] Similarly, Adolph Eduard Grube advanced the taxonomy around 1850 by describing multiple Nereis species and emphasizing paragnath arrangements on the proboscis as diagnostic features, aiding in distinguishing Nereis from related genera.^[10] These efforts were followed by Anders Johan Malmgren's 1865 revisions, which introduced subgenera such as Eunereis and Hediste to address morphological variability within Nereis, signaling the onset of systematic splits in the 19th century.^[11] Throughout the 20th century, ongoing taxonomic revisions further fragmented the original Nereis concept, with many species reassigned to new genera like Neanthes (Kinberg, 1865) and Alitta (Kinberg, 1865) based on detailed morphological studies of setal patterns and reproductive structures.^[7] Post-2000 phylogenetic analyses, incorporating DNA barcoding of the COI gene, have continued to refine these boundaries by revealing cryptic diversity and non-monophyly in traditional Nereis groupings, as demonstrated in studies of North Atlantic and global nereidid populations.^[12] For instance, molecular data have supported the elevation of subgenera to full genera and identified new species complexes, enhancing the precision of Nereis classification in contemporary systematics.^[13]

Morphology and Anatomy

External Morphology

Nereis species possess an elongated, segmented, tube-like body that is bilaterally symmetrical and slightly dorsoventrally flattened, divided into three main regions: the prostomium (head), a trunk of numerous metameres (up to 200 segments), and the pygidium (tail). The body length typically ranges from 5 to 30 cm, though some individuals can exceed 1 meter. The integument is covered by a thin, iridescent collagenous cuticle featuring microscopic striations that produce colors ranging from green to brown, often with a visible dorsal blood vessel appearing as a dark median line in live specimens.^[14]^[15]^[16] The prostomium is a small, flattened, pear-shaped plate at the anterior end, equipped with two pairs of black or reddish eyes arranged in a trapezoid, two short antennae for sensory perception, and two large fleshy palps divided into proximal and distal articles. Immediately posterior to the prostomium lies the peristomium, an apodous ring encircling the mouth, which bears four pairs of long, tentacular cirri. Each trunk segment, except the first and last, features a pair of fleshy parapodia—lateral appendages that are biramous, consisting of a dorsal notopodium and a ventral neuropodium; these are adorned with ligules, cirri, and bundles of chitinous chaetae, including compound notosetae and neurosetae supported by acicula rods. The pygidium forms a simple posterior tagma with the anus and two long anal cirri derived from modified parapodia.^[14]^[15]^[16] Morphological variations occur across life stages, particularly between the atokous (non-reproductive, benthic) and epitokous (reproductive, pelagic) forms. In the atokous form, parapodia are compact and adapted for crawling, with standard chaetae facilitating burrowing and substrate adhesion. The epitokous form, developed during maturation, exhibits modifications for swarming, including enlarged eyes, enhanced nuchal organs, and transformed parapodia bearing paddle-like swimming setae that replace the typical compound chaetae, enabling rapid vertical migration in water columns. These external adaptations underscore the dimorphic life strategy in Nereis, optimizing survival in intertidal and subtidal environments.^[14]^[15]

Internal Anatomy

The digestive system of Nereis consists of a complete, straight tubular alimentary canal extending from the mouth to the anus, divided into foregut, midgut, and hindgut regions. The foregut includes the buccal cavity, a muscular pharynx that can be everted for feeding, and an esophagus; the pharynx is equipped with a proboscis armed with paragnaths, which are chitinous, tooth-like structures aiding in prey capture.^[14] The midgut comprises the intestine, lined with glandular epithelium that secretes digestive enzymes, while the hindgut features a short rectum leading to the anus.^[17] This system facilitates extracellular digestion of organic matter, with absorption primarily occurring in the intestine.^[18] The circulatory system is closed and segmental, featuring a dorsal longitudinal vessel that acts as the main pumping structure through peristaltic contractions, propelling blood anteriorly, and a ventral vessel that returns blood posteriorly.^[14] Segmental vessels connect these longitudinal channels to capillary beds in the parapodia, gut, and body wall, enabling nutrient and oxygen distribution; additional contractile "hearts" in certain segments enhance circulation.^[19] The excretory system comprises paired metanephridia in each body segment (except the first and last), with a ciliated nephrostome opening into the coelom for filtration, a convoluted duct for reabsorption, and an external nephridiopore for ammonia excretion, also serving osmoregulatory functions in marine environments.^[14] The nervous system is centralized yet segmental, with a dorsal supraesophageal brain located in the prostomium, connected via circumpharyngeal connectives to a subesophageal ganglion and a double ventral nerve cord running posteriorly.^[14] The ventral cord features fused segmental ganglia that control local reflexes, linked by intersegmental connectives; giant fibers in the cord facilitate rapid escape responses.^[20] Respiratory functions rely on cutaneous gas exchange across the thin body wall and through vascularized parapodial structures, particularly the dorsal ligules of the notopodia, which increase surface area for oxygen uptake in aquatic habitats.^[14] Sensory structures include paired nuchal organs on the posterior prostomium, which are ciliated chemoreceptors detecting environmental chemicals and aiding in food location.^[21] Statocysts, small sac-like organs in the prostomium, provide equilibrium sensing via statoliths that stimulate hair cells during movement.^[21] These adaptations, combined with palps and antennae, support sensory integration for navigation and predation.^[14]

Life Cycle and Reproduction

Reproductive Strategies

Nereis species primarily reproduce sexually, with no evidence of parthenogenesis or other forms of asexual reproduction such as fission, though they exhibit regenerative capabilities for somatic tissue repair.^[22] These polychaete worms are gonochoristic, meaning they are dioecious with separate sexes.^[23] Gametes develop within the coelom or its derivatives, maturing in specialized coelomic clusters before release.^[14] A key aspect of the reproductive biology of many Nereis species is the heteronereid transformation, where immature atokous forms—benthic, non-swimming individuals adapted for feeding and burrowing—undergo metamorphosis into epitokous forms optimized for reproduction. This process involves hormonal cues that trigger the development of modified posterior segments, including hypertrophied parapodia for swimming and enlarged gonads filled with gametes, while anterior segments remain atokous for sensory functions. The transformation results in a distinct border between atokous and epitokous regions, enabling the worm to leave its habitat and participate in pelagic swarming; however, reproductive strategies vary across the genus, with some species reproducing iteratively without such transformation.^[24] Gamete production occurs in the epitokous phase, with females developing large oocytes that accumulate yolk in the coelomic cavity and males producing spermatocytes in similar clusters, often concentrated in the posterior body. Fertilization is external, with gametes broadcast into the water column during spawning events to maximize encounter rates in dilute concentrations.^[25] Mating in Nereis involves synchronized swarms of epitokes, often timed to lunar cycles—such as four days after the full moon in N. virens—to coincide with optimal tidal and light conditions for dispersal. During these nuptial dances, males and females swim rapidly in the water column, releasing sperm and eggs simultaneously; in some species, brief pseudocopulation-like interactions may facilitate closer gamete transfer, though broadcast spawning predominates.^[26] These swarms are semelparous, with individuals typically dying after a single reproductive event.^[24] Fecundity varies by species and body size, with females capable of producing thousands to over a million eggs per spawning; for instance, N. virens individuals up to 54 cm long release up to 1.3 million eggs, each approximately 180–200 µm in diameter.^[26] This high output supports the production of numerous offspring to offset mortality in the planktonic phase.^[25]

Developmental Stages

Development in the genus Nereis begins with external fertilization, typically occurring during epitokous swarming events where mature adults release gametes into the water column. The zygote undergoes spiral cleavage, a characteristic pattern in polychaete annelids where early cell divisions produce a stereotypic arrangement of blastomeres in oblique, alternating directions, leading to the formation of the trochophore larva.^[27] The trochophore larva is a free-swimming, ciliated stage featuring a spherical body with a prototroch (ciliary band for locomotion and feeding) and an emerging stomodaeum. This stage progresses through early, middle, and late phases, during which the hyposphere elongates, the telotroch forms, and initial segmentation begins, enabling the larva to capture planktonic food particles.^[27]^[28] Following the trochophore, the larva transitions to the metatrochophore and nectochaete stages, marked by the development of body segments and setae. The nectochaete possesses functional parapodia, chaetae for locomotion, and a complete digestive system, allowing active swimming and feeding near the substrate; this phase precedes metamorphosis.^[27] Metamorphosis transforms the nectochaete into a juvenile worm capable of benthic life; in some species, this involves settlement to the substrate and initiation of tube-building or burrowing behaviors. The process includes loss of larval cilia, elongation of the body, and formation of additional segments, culminating in the young worm resembling the adult form.^[27] Environmental factors significantly influence survival and developmental success. Salinity affects fertilization, with optimal rates (≥70–75%) occurring between 22–34‰, while trochophore and nectochaete stages tolerate a broader range of 14–45‰ due to developing osmoregulatory structures like protonephridia. Temperature accelerates development and metamorphosis, with rates increasing from 5°C to 23°C, the highest survival and transformation observed at warmer temperatures (17–23°C) above 14‰ salinity, though extreme values reduce viability.^[29]

Ecology and Distribution

Habitats and Geographic Range

Nereis species exhibit a cosmopolitan distribution, occurring in marine waters across temperate and tropical regions worldwide, with presence reported in all major oceans.^[1] They are particularly abundant in coastal areas of the North Atlantic, Pacific, and European shorelines, reflecting their broad ecological adaptability.^[30] These polychaetes primarily inhabit soft-bottom environments such as intertidal zones, subtidal sediments, and estuaries, where they construct temporary burrows in mud, sand, or under rocks.^[11] They are common in soft-sediment communities from the intertidal zone to abyssal depths, contributing to benthic ecosystems in these dynamic settings.^[31] Some species demonstrate euryhaline capabilities, tolerating low salinities in brackish or even ephemeral freshwater habitats, as exemplified by N. diversicolor.^[32] Nereis worms have evolved physiological and behavioral adaptations to thrive in variable conditions, including tolerance to low oxygen levels through hypoxia-resistant mechanisms and osmotic regulation in fluctuating salinities.^[33] Burrow ventilation is achieved via peristaltic body movements, which facilitate water flow to maintain oxygen supply and remove wastes in oxygen-poor sediments.^[34]

Feeding, Behavior, and Interactions

Nereis species are predominantly carnivorous or omnivorous predators and scavengers, consuming small invertebrates such as crustaceans, gastropods, and polychaetes, along with detritus, diatoms, and algae. They capture prey by everting the pharynx, which rapidly extends to expose chitinous jaws and denticles that grasp and sever food items before retraction into the mouth. ^[14] ^[35] Within their burrows, individuals scavenge organic matter and opportunistic prey, with feeding activity influenced by environmental factors like tides and storms. ^[36] As errant polychaetes, Nereis exhibit a mobile lifestyle, crawling across sediments or prospecting nocturnally using parapodia for undulating locomotion. ^[14] ^[37] Activity patterns follow circadian, circatidal, and circalunidian rhythms, with peaks during low tide or darkness under free-running conditions, enabling synchronization with environmental cues for foraging and predator avoidance. ^[38] Escape responses to predators include rapid burrowing withdrawal triggered by chemical alarm cues from damaged conspecifics and vigorous body thrashing to dislodge attackers. ^[39] In marine communities, Nereis play key ecological roles as both predators and prey. They prey on juvenile bivalves like Macoma balthica and other infauna, reducing densities and structuring benthic assemblages through direct consumption and disturbance. ^[40] ^[41] As common forage, they support populations of fish such as flounder and wading birds in estuaries, transferring energy up the food web. ^[42] Bioturbation via burrow construction and sediment reworking enhances nutrient cycling, increasing oxygen penetration and organic decomposition rates in intertidal mudflats. ^[43] Certain species, including N. fucata, form commensal associations with hermit crabs, residing within occupied shells for protection while benefiting the host minimally. ^[44]

Diversity and Species

Species Diversity

The genus Nereis encompasses approximately 226 species attributed to it, many subject to ongoing taxonomic revisions that refine species boundaries within the group.^[3] The broader family Nereididae demonstrates substantial diversity, comprising over 700 species distributed across 45 genera, underscoring the phylogenetic richness of this polychaete lineage.^[9]^[11] Species diversity in Nereis is most pronounced in coastal environments of the Indo-Pacific and Atlantic regions, where environmental heterogeneity supports varied ecological niches.^[45] Genetic analyses have increasingly uncovered cryptic species complexes, revealing morphologically indistinguishable lineages that were previously overlooked through traditional approaches. These findings emphasize the Indo-Pacific as a hotspot for hidden diversity, driven by molecular markers like COI and 16S rRNA genes.^[45] Recent additions include Nereis quadrata from the Persian Gulf, described in 2025.^[46] Challenges in species identification arise from the limitations of morphological taxonomy, which often fails to distinguish subtle variations, contrasted with the precision offered by molecular methods that detect genetic divergence. Certain populations face threats from anthropogenic pollution and habitat degradation in urbanized coastal zones.^[47] For example, Nereis virens serves as a representative species commonly found in temperate Atlantic intertidal habitats.^[48]

Notable Species and Synonyms

Nereis succinea, now classified as Alitta succinea (Leuckart, 1847), is a prominent species within the former Nereis genus, serving as the type species for the genus Alitta. It is commonly found in estuarine environments, where it inhabits soft sediments and exhibits distinctive red pigmentation on the dorsal surface, often extending from the head and anterior parapodia. This coloration, combined with a darkly pigmented head and pale red or greenish-yellow posterior region dotted with white or dark spots, aids in its identification. Synonyms for A. succinea include Nereis limbata Ehlers, 1868, considered a subjective synonym based on morphological similarities in type material from the North Sea and American coasts.^[49]^[50]^[51] Nereis virens, reclassified as Alitta virens (Sars, 1835), is one of the largest nereidid species, capable of reaching lengths up to 90 cm, though typically measuring 20-40 cm in adulthood. It displays a characteristic green coloration in males and an orange-green metallic hue in females, inhabiting sandy or muddy substrates in the North Atlantic from Virginia northward to Arctic regions. This species is widely harvested as commercial fishing bait due to its robust size and active swimming behavior, supporting recreational and commercial fisheries in areas like the Gulf of Maine.^[52]^[14]^[53]^[26] Nereis diversicolor, currently recognized as Hediste diversicolor (O.F. Müller, 1776), is a euryhaline polychaete adapted to a wide salinity range, commonly burrowing in brackish marshes and estuarine mudflats. It reaches lengths of 6-12 cm and plays a key role as an ecological indicator species, particularly for organic pollution and heavy metal contamination, due to its sensitivity and bioaccumulation properties in polluted sediments. Its burrowing activity enhances sediment oxygenation and nutrient cycling in low-salinity habitats.^[54]^[55]^[56]^[57] Taxonomic revisions have led to the reclassification of many species formerly under Nereis, reflecting phylogenetic distinctions in chaetal morphology and reproductive traits. For instance, Nereis japonica Izuka, 1908, is now placed in Neanthes japonica, based on the absence of homogomph falcigers in posterior notopodia. Similarly, N. diversicolor has been transferred to Hediste Malmgren, 1867, due to fused falcigers in the supra-acicular neuropodial setae, with ongoing studies identifying cryptic lineages within this complex. These shifts highlight the polyphyletic nature of the original Nereis genus.^[58]^[59]^[54]^[60] Species within the Nereis complex hold economic significance, particularly as fishing bait and in aquaculture. A. virens and H. diversicolor are intensively collected or cultured for bait in recreational fisheries across the Atlantic, with A. virens pioneering intensive aquaculture systems since the 1980s to meet demand and reduce wild harvest pressures. In integrated systems, these polychaetes process fish farm effluents, providing high-protein feed while improving waste management sustainability.^[61]^[62]^[63]^[64]

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Jul 9, 2025 · Nereididae is a prolific annelid family widely distributed in the world oceans, especially in the Indo-Pacific Convergence Zone (IPCZ).
[47]
(PDF) Intertidal Polychaetes as Indicators of Pollution Resulting from ...
May 17, 2021 · The study is aimed at identifying the intertidal polychaete species and their abundance in these areas, and using them as indicators of pollution.
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https://www.marinespecies.org/aphia.php?p=taxdetails&id=129433
[49]
http://marinespecies.org/aphia.php?p=taxdetails&id=234850
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Alitta succinea (Leuckart, 1847) - GBIF
red or reddish brown on the dorsal surface, which often extends from the head and parapodia of the anterior segments. ” Preserved coloration. Type material has ...
[51]
https://www.marinespecies.org/aphia.php?p=taxdetails&id=339377
[52]
http://marinespecies.org/aphia.php?p=taxdetails&id=129464
[53]
Sandworm (Nereis) — Live Science Specimen - Gulf Of Maine, Inc.
In stockMales are commonly green and brown in color while the larger females are orange green metallic colors. ... bait in mid-Atlantic markets. Diggers can get as many ...
[54]
http://marinespecies.org/aphia.php?p=taxdetails&id=152302
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Ragworm (Hediste diversicolor) - The Marine Life Information Network
Colour varies (see general biology) but mainly green or reddish. Prominent dorsal blood vessel. Inhabits mucous burrow. Additional information. The form and ...Missing: color | Show results with:color
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Response of Nereis diversicolor population (Polychaeta, Nereididae ...
Jan 20, 2008 · The ragworm Nereis diversicolor O.F. Müller, 1776 is a brackish and euryhaline species and an indicator of organic pollution (Scaps and Borot, ...
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Biomarkers in <i>Nereis diversicolor</i> (Polychaeta - Scientia Marina
Mar 30, 2013 · Biomarker responses of N. diversicolor are useful tools for environmental quality assessment on the southwest Iberian coast.
[58]
http://marinespecies.org/aphia.php?p=taxdetails&id=334096
[59]
A review of Asian Hediste species complex (Nereididae, Polychaeta ...
Aug 10, 2025 · A review of Asian Hediste species complex (Nereididae, Polychaeta) with descriptions of two new species and a redescription of Hediste japonica ...Missing: reclassified | Show results with:reclassified
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The curious and intricate case of the European Hediste diversicolor ...
Oct 17, 2022 · Past molecular studies using mtDNA sequences and alloenzymes signalled the existence of at least two cryptic species within the Hediste ...
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Nereis virens (sand worm) | CABI Compendium
Jan 4, 2023 · This datasheet on Nereis virens covers Identity, Overview, Distribution, Biology & Ecology, Environmental Requirements, Uses, Management ...
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[PDF] Bait worms: a valuable and important fishery with implications for ...
At local, regional and national scales, polychaete bait fisheries are highly valuable, extract significant biomass and have considerable impacts; therefore, ...
[63]
Polychaete aquaculture and polychaete science: a mutual synergism
In 1984, the commercial aquaculture of the polychaete Nereis virens was initiated in NE England, at about the same time similar developments took place in.
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Utilization of waste from a marine recirculating fish culture system as ...
Dec 21, 2011 · This species is an excellent candidate for integrated aquaculture and waste recycling. Highlights. ▻ The polychaete worm Nereis virens was fed ...