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Mysida

Mysida, commonly known as mysid shrimps or opossum shrimps, are a diverse order of small, shrimp-like crustaceans within the superorder Peracarida of the class Malacostraca.^[1] They are distinguished by their caridoid facies, featuring a carapace that envelops most of the thorax, movable stalked eyes (which may be reduced in some species), and a brood pouch or marsupium in females for embryonic development.^[2] Typically ranging from 5 to 25 mm in length, mysids encompass two families—Mysidae and Petalophthalmidae—with 183 genera and 1,233 described species (as of 2025).^[3] These crustaceans exhibit a broad ecological distribution, inhabiting freshwater, brackish, coastal, pelagic, and deep-sea environments across all continents from 80°N to 80°S latitude.^[4] Many species are epi- or hyperbenthic, while others are fully pelagic or even subterranean, demonstrating remarkable adaptability to varied salinities, temperatures, and oxygen levels.^[2] Biologically, mysids are primarily omnivorous filter feeders, consuming phytoplankton, zooplankton, detritus, and meiofauna, which positions them as key intermediaries in aquatic food webs.^[4] Reproduction occurs via direct development in the female's marsupium, where embryos hatch as juveniles before release, often synchronized with the mother's molt cycle; fecundity varies by species but supports rapid population growth in favorable conditions.^[4] Notable features include statocysts in the uropods for balance (in Mysidae) and swarming behaviors that enhance predator avoidance and nutrient cycling.^[2] Their ecological significance extends to serving as primary prey for fish, birds, mammals, and other invertebrates, thereby linking benthic and pelagic realms and influencing community structure in estuarine and marine ecosystems.^[4]

Description and Anatomy

Physical Characteristics

Mysids exhibit a distinctive shrimp-like body plan, characteristic of the caridoid facies typical of many peracarid crustaceans, with the body divided into three main regions: a cephalon, thorax, and abdomen. The carapace forms a dorsal shield that envelops most of the thorax and is fused with the first four thoracic somites, leaving the posterior thoracic segments and abdomen exposed for flexibility. This structure typically measures 1–30 mm in length, though some deep-water species can reach up to 80 mm.^[2]^[5] A key distinguishing feature in females is the presence of a ventral brood pouch, or marsupium, formed by oostegites on the thoracic appendages, which gives mysids their common name "opossum shrimps." The cephalon bears stalked compound eyes and paired antennules and antennae; the antennules are three-segmented with flagella that serve sensory functions, while the antennae feature a scale-like exopod armed with plumose setae for both sensory detection and manipulation. The eight thoracic appendages include the first two pairs modified as maxillipeds for handling food and grooming, with the remaining pairs functioning as walking legs.^[2]^[5]^[6] The abdomen consists of six somites, with biramous natatory pleopods that are often reduced in females but more developed in males for swimming. Terminally, the telson—a flattened, elongate plate with serrated or setose margins and a cleft or entire apex armed with spines—articulates with the uropods to form a fan-like tail structure essential for rapid backward propulsion. Across species, body size varies from under 5 mm in some freshwater forms to over 20 mm in marine ones, and coloration ranges from translucent in pelagic species, aiding camouflage in open water, to more opaque or pigmented in benthic types.^[2]^[5]

Internal Anatomy

The internal anatomy of Mysida reveals a suite of organ systems adapted for their primarily pelagic or benthic lifestyles in marine and estuarine environments, with physiological features that support efficient nutrient processing, circulation, gas exchange, neural coordination, and osmoregulation. As members of the Peracarida, Mysida exhibit variations in internal structures compared to other crustacean groups, such as a relatively simplified arterial network and specialized filtering mechanisms in the gut that reflect their opportunistic feeding habits.^[7]^[8] The digestive system comprises a foregut, midgut, and hindgut, facilitating mechanical breakdown, enzymatic digestion, and waste elimination. The foregut includes a cardiac stomach equipped with a gastric mill featuring dorsal, lateral, and ventral chitinous ridges that triturate food particles, while the pyloric stomach is divided into chambers by ridges and stiff hairs to filter coarse material from fluids and fine particulates.^[9] The midgut gland, or hepatopancreas, consists of five paired lobes at the stomach-midgut junction, where it secretes digestive enzymes and absorbs nutrients from processed fluids, differing from the more extensive glandular arrays in decapods by its fewer but larger caeca suited to Peracarida diets.^[9]^[7] Coarse residues pass to the hindgut for compaction and expulsion through the anus, with the overall system emphasizing filtration over extensive grinding seen in herbivorous crustaceans.^[7] Mysida possess an open circulatory system typical of malacostracans, with a tubular heart located in the thorax that pumps hemolymph into tissues via arteries before it returns through open sinuses. The heart, extending through much of the thoracic region, features an anterior aorta branching into four pairs of arteries supplying the head, antennae, and eyes, along with fewer lateral cardiac arteries (typically five pairs) compared to the up to ten pairs in related Lophogastrida, reflecting a less complex vascularization adapted to their smaller body sizes.^[8] A posterior aorta distributes hemolymph to the abdomen, and myoarterial formations with internalized muscles in the cephalothorax aid propulsion, a peracarid-specific trait that enhances efficiency over the more rigid arterial systems in eumalacostracans like decapods.^[8]^[10] Respiration in Mysida occurs through diffusion across the thin carapace and body surface, as they lack gills. Oxygen uptake is facilitated by the branchial chamber formed by the carapace and appendage beating, with adaptations enabling efficient uptake in low-oxygen habitats.^[11] This system emphasizes passive diffusion augmented by locomotion, contrasting with the gill-based ventilation in groups like decapods. The nervous system centers on a supraesophageal ganglion, or brain, located in the head and comprising protocerebral, deutocerebral, and tritocerebral regions, with prominent deutocerebral olfactory neuropils linked by globular tracts for chemosensory processing. This brain connects anteriorly to compound eyes on movable eyestalks in epigean (surface-dwelling) species, featuring layered visual neuropils like the lamina and medulla for image formation, while a ventral nerve cord runs posteriorly with segmental ganglia fusing thoracic and abdominal neuromeres for motor control.^[12] Unlike the more centralized brains in higher crustaceans, the Mysida system integrates sensory inputs from antennae briefly, supporting rapid responses to environmental cues without extensive fusion of ganglia.^[12] Excretion and osmoregulation are handled by paired green glands, or antennal glands, situated at the base of the antennae within the antennal sympods, which filter hemolymph to remove nitrogenous wastes and regulate ion balance in fluctuating salinities. Each gland includes a coelomic sac, labyrinth for reabsorption, and bladder opening near the antennal base, enabling hyperosmotic urine production crucial for estuarine species facing salinity gradients.^[13]^[14] This setup mirrors other malacostracans but is particularly vital in Mysida for maintaining hemolymph osmolarity during vertical migrations or euryhaline transitions, with the end sac facilitating active ion transport.^[14]^[13]

Habitat and Distribution

Global Range

Mysida display a cosmopolitan distribution, primarily as marine organisms inhabiting all major oceans worldwide, including the Atlantic, Pacific, Indian, Arctic, and Southern Oceans.^[15] Their presence extends to freshwater systems globally, where approximately 72 species occur, representing about 6.7% of the total Mysida diversity and often as glacial relicts or euryhaline invaders from estuarine origins.^[16] Brackish environments, such as estuaries and coastal lagoons, also support diverse Mysida populations across continents.^[15] In terms of depth, Mysida occupy a broad vertical range from shallow coastal waters to abyssal depths exceeding 4,000 meters, with some families like Petalophthalmidae restricted to deep-sea habitats.^[15] Zonation patterns vary by species and region; for instance, certain pelagic forms aggregate in epipelagic swarms within open ocean waters, facilitating energy transfer in the water column, while benthic species predominate in coastal sediments.^[17]^[18] Subterranean populations further highlight their adaptability, with cave-dwelling species recorded in anchialine and freshwater aquifers of Europe, including the Adriatic karst systems, and North America, such as cenotes in Mexico and blue holes in the Bahamas.^[15] Endemic hotspots concentrate in ancient lake systems like the Ponto-Caspian basin, home to 24 species, approximately 20 of which are endemic primarily within the genus Paramysis, underscoring a significant radiation event.^[16] Human-mediated invasions have expanded their ranges, exemplified by Hemimysis anomala, a Ponto-Caspian native now established in the Laurentian Great Lakes since 2006, where it coexists with the native Mysis diluviana.

Environmental Preferences

Mysids exhibit a wide range of salinity tolerances, spanning from euhaline marine conditions of 30-35 parts per thousand (ppt) to oligohaline freshwater environments below 5 ppt, enabling their presence in diverse aquatic systems worldwide. Euryhaline species, such as Neomysis integer and Neomysis mercedis, thrive in fluctuating estuarine habitats, tolerating salinities from 0.5 to over 25 ppt, which facilitates their adaptation to brackish zones where salinity gradients are pronounced.^[4]^[19] In contrast, strictly marine species like Americamysis bahia prefer salinities above 15 ppt, with optimal growth occurring between 20 and 34 ppt, highlighting how salinity preferences shape their ecological niches in coastal and open-water settings.^[4] Temperature preferences among mysids vary significantly with geographic distribution, from polar species enduring cold waters below 5°C to tropical forms active in environments up to 30°C. For instance, the polar mysid Mysis relicta performs optimally in temperatures of 4-15°C during its diel vertical migrations, reflecting adaptations to stable, low-temperature regimes in high-latitude lakes and seas.^[20] Tropical species, such as those in Malaysian mangrove estuaries like Mesopodopsis orientalis, tolerate warmer conditions around 25-30°C, demonstrating the order's broad thermal plasticity across latitudinal gradients.^[21] Eurythermal coastal mysids, including Neomysis americana, can withstand ranges from 6 to 34°C, though sublethal effects on growth and survival intensify near upper limits.^[22] Substrate preferences in mysids include pelagic lifestyles in the water column, benthic associations with mud or sand, and interstitial habitation within sediments, allowing them to exploit varied microhabitats. Pelagic species, such as certain Mysis forms, remain free-swimming in mid-water layers, while many others are hyperbenthic, crawling or burrowing into soft sediments like sand or mud for refuge.^[23] Benthic mysids, exemplified by Gastrosaccus psammodytes, favor substrates with specific particle sizes that facilitate burrowing, often in coastal or estuarine sands.^[4] Interstitial species inhabit the pore spaces between gravel or coral sediments, particularly in cryptic environments, enhancing their survival in dynamic benthic zones.^[23] Mysids generally prefer normoxic waters with dissolved oxygen levels above 4-5 mg/L, though some coastal species demonstrate notable hypoxia tolerance down to 1.5 mg/L, enabling persistence in seasonally deoxygenated habitats. For example, Tenagomysis novae-zealandiae maintains activity at 1.5 mg O₂/L but experiences high mortality below 0.5 mg O₂/L, underscoring their sensitivity to severe anoxia.^[24] Deep-sea mysids are adapted to low-oxygen environments in oxygen minimum zones while remaining aerobic. This tolerance varies by species, with estuarine forms showing behavioral adjustments to moderate hypoxia but avoiding prolonged exposure.^[4] Many mysids display photophobic behavior, avoiding bright light and favoring nocturnal or deep-water activity to minimize predation risk, which influences their vertical distribution in the water column. Species such as Mysis diluviana exhibit extreme photophobia, rapidly fleeing even low light levels during daytime, often retreating to benthic or profundal zones.^[25] This negative phototaxis, observed in taxa like Hemimysis lamornae, alternates with weak positive responses in low-light conditions, promoting crepuscular feeding.^[26] Light avoidance is particularly pronounced in freshwater and coastal mysids, linking their activity patterns to diurnal cycles and water depth.^[27]

Biology and Behavior

Locomotion and Sensory Systems

Mysids exhibit diverse modes of locomotion adapted to their benthic, epibenthic, or pelagic lifestyles. Swimming is the primary form of movement for many species, achieved through the coordinated beating of biramous pleopods on the abdomen, which provide sustained propulsion, supplemented by the exopods of thoracic pereopods for additional thrust.^[28] The uropods and telson form a tail fan that enables rapid backward escapes via powerful flexion of the abdomen, a common antipredator response in shrimp-like crustaceans. Benthic and epibenthic mysids often crawl along substrates using their thoracic pereopods, which bear setae for traction on mud, sand, or algae. Some species, particularly in the subfamily Gastrosaccinae, are specialized burrowers, employing modified pereopods to excavate shallow tunnels in soft sediments for refuge during the day, emerging nocturnally to feed. Sensory systems in mysids facilitate navigation, predator avoidance, and resource detection in varied environments. Statocysts located on the endopods of the uropods serve as balance organs, detecting gravity and angular acceleration to maintain orientation during swimming or burrowing. Chemoreceptors, primarily aesthetasc sensilla on the antennules, detect chemical cues such as food odors or pheromones, aiding in foraging and mate location.^[29] Mechanoreceptors, including setal arrays on the antennae and body, sense water currents and vibrations, enabling responses to hydrodynamic disturbances from nearby organisms.^[30] Vision varies markedly across habitats; pelagic species possess stalked compound eyes sensitive to low light levels for detecting silhouettes or bioluminescent signals, with spectral sensitivities peaking in the blue-green range to match oceanic light penetration.^[31] In contrast, cave-dwelling or deep-sea forms, such as those in the family Petalophthalmidae, often lack functional eyes or have reduced, non-pigmented ocular structures, relying instead on enhanced chemosensory and mechanosensory inputs.

Feeding Mechanisms

Mysids employ a variety of feeding mechanisms adapted to their diverse habitats, primarily consisting of filter-feeding, raptorial predation, and scavenging. In pelagic species, filter-feeding predominates, where fine setae on the endopods of thoracic appendages (particularly the second through eighth pereopods) create a straining basket to capture suspended particles such as plankton and detritus from the water column as the mysids swim or ventilate water over their gills.^[32] This mechanism allows efficient collection of small food items, with the setae acting as a sieve that directs particles toward the mouthparts for ingestion.^[33] For larger prey, mysids utilize raptorial feeding, employing modified gnathopods and other thoracic appendages to grasp and manipulate small invertebrates, algae, or other suitable items. This active predation involves rapid extension of the endopods to seize food, often targeting motile prey like copepods or rotifers, and is particularly common in more mobile or coastal species such as Paramesopodopsis rufa and Neomysis integer.^[33] In benthic or semi-benthic environments, scavenging plays a key role, with mysids consuming organic matter from the substrate; here, robust mandibles equipped with grinding teeth break down tougher particles like detritus or carrion before swallowing.^[33] The diet of mysids reflects their opportunistic omnivory, encompassing primarily zooplankton (e.g., copepods, cladocerans, rotifers), phytoplankton (e.g., diatoms), bacteria, and detritus, with preferences shifting based on availability and size—smaller individuals favoring finer particles while larger ones incorporate more animal matter.^[33] Species like Neomysis mercedis selectively target large diatoms and copepods, while Leptomysis spp. rely heavily on sedimentary organic matter and algae.^[33] This flexibility enables mysids to exploit varied resources, including terrestrial inputs like plant fragments in coastal areas.^[33] Daily rations in mysids vary widely by species, prey type, and environmental conditions, typically ranging from 3% to 30% of body carbon weight, though higher rates up to 200% have been recorded under optimal feeding scenarios such as continuous access to cladocerans.^[33] For instance, Neomysis americana can consume over 100% of its body carbon daily when feeding on Artemia nauplii, highlighting the influence of food density and quality on intake.^[34] These consumption levels support their high metabolic demands and reproductive output.^[33]

Reproduction and Life History

Reproductive Strategies

Mysids primarily engage in sexual reproduction characterized by internal fertilization, where males transfer spermatophores containing sperm to females using modified thoracopods, typically the eighth pair, which function as genital papillae or lobes. This transfer often occurs rapidly, within minutes after the female molts, when the empty marsupium is accessible, allowing sperm to be deposited directly into the brood pouch for subsequent egg fertilization.^[35]^[36] Following fertilization, females brood the embryos in a ventral marsupium formed by oostegites on the thoracopods, which provides mechanical protection against predation and environmental stressors while facilitating oxygenation through water currents generated by the female's appendages and adjacent gills. The number of oostegites varies by family, ranging from 2–3 pairs in Mysidae to 7 pairs in groups like Petalophthalmidae, enabling effective enclosure of developing young. This brooding strategy ensures direct development, with juveniles emerging fully formed after several embryonic and post-embryonic stages.^[35]^[15] The vast majority of mysid species are dioecious, with distinct males and females exhibiting sexual dimorphism, such as larger female body sizes and modified male appendages for sperm transfer; hermaphroditism is exceedingly rare and not well-documented in the order. Breeding patterns typically feature seasonal peaks tied to environmental cues like temperature, photoperiod, and food abundance, with temperate species often producing multiple generations annually (bivoltine to trivoltine cycles) and continuous reproduction in some tropical or subtropical populations.^[35]^[37] Fecundity in mysids generally ranges from 10 to 40 eggs per brood, though this varies with female body size, species-specific traits, and habitat conditions, with larger individuals in nutrient-rich environments producing more eggs. Mating events are brief and nocturnal, often involving males attending receptive females post-molt, potentially guided by chemical cues, but elaborate courtship displays are not prominently reported.^[38]^[36]^[37]

Developmental Stages

Mysids exhibit epimorphic development, in which embryos develop directly within the female's marsupium (brood pouch) and hatch as miniature adults lacking a distinct larval phase.^[39]^[40] Embryonic development typically spans 2-4 weeks, depending on species and environmental conditions; for instance, in Metamysidopsis elongata, it lasts approximately 20-24 days at 21-23°C, progressing through stages including prenaupliar, nauplioid, and postnauplioid phases marked by egg membrane shedding and appendage formation.^[40] In Americamysis bahia, the initial embryonic phase alone requires 1.5-3.6 days across temperatures of 16-29°C, with full marsupial development completing prior to juvenile release.^[41] Juveniles emerge from the marsupium fully formed, resembling adults in morphology but smaller in size, typically measuring 1-1.3 mm in length at release.^[39]^[40] Post-release, juveniles undergo multiple molts (typically 5-10) during gradual growth to adulthood, with no intervening larval stages. In Metamysidopsis insularis, juveniles experience about 8 molts over the first 16 days post-release at 25°C, transitioning through early (0-2 days, 1 molt), mid (3-6 days, 3 molts), and late (7-12 days, 3 molts) juvenile phases, resulting in size increases from ~1.1 mm to nearly 4 mm.^[39] Growth rates are strongly influenced by temperature and food availability; higher temperatures shorten intermolt periods and accelerate overall growth without altering molt increments, while optimal feeding (e.g., frequent Artemia nauplii) enhances size gains, as observed in estuarine species like Tenagomysis chiltoni where elevated temperatures boosted growth but reduced survival.^[42]^[43] Most mysid species have lifespans of 1-3 years under natural conditions, though laboratory-cultured individuals like M. insularis may live only ~3 months; longer durations, such as 2.5 years in Mysis oculata, occur in colder boreal habitats.^[39] Sexual maturity is attained after 3-6 months in many species, varying with temperature and habitat; in warmer conditions, it occurs in as little as 10-17 days (e.g., summer cohorts), but extends to 57-86 days in colder seasons.^[44] Reproductive strategies differ by habitat, with semelparity (single breeding event followed by death) common in boreal or high-latitude species like Mysis gaspensis, and iteroparity (multiple broods) prevalent in temperate estuarine populations such as Neomysis americana, where overwintering females produce successive cohorts over 8-10 months.^[45]^[46] In females, the molting process (ecdysis) is synchronized with reproduction to minimize brood loss; young are released just prior to ecdysis, followed by copulation and egg deposition into the newly formed marsupium, as seen in the 12-day molt-breeding cycle of Siriella armata.^[47]^[48] This coordination ensures ovarian development, marsupial formation, and brood maturation align across the female's integumental cycle.^[4]

Ecology and Interactions

Trophic Role

Mysids occupy a pivotal position in aquatic food webs as primary and secondary consumers, primarily functioning as planktivores that link phytoplankton and detritus to higher trophic levels. By grazing on microalgae, zooplankton such as copepods and cladocerans, and organic detritus, they efficiently transfer energy from basal producers to predatory invertebrates and fish, enhancing overall ecosystem productivity.^[49]^[50]^[51] This role is amplified in their feeding habits, which emphasize selective planktivory in pelagic environments.^[51] Their substantial biomass contributions further underscore this trophic linkage, with swarms achieving high densities reaching thousands of individuals per cubic meter in coastal and estuarine systems, indirectly supporting fisheries through sustained prey availability for commercially important species.^[52] These aggregations not only concentrate energy but also facilitate rapid biomass turnover, bolstering the resilience of food webs against perturbations.^[53] Mysids contribute to nutrient cycling via diel vertical migrations, where populations ascend to surface waters at night to feed and descend to benthic zones by day, transporting organic matter, nutrients, and carbon between pelagic and benthic habitats. This migration promotes nutrient recycling through excretion and fecal pellet deposition, influencing primary production and maintaining biogeochemical balances in stratified lakes and coastal areas.^[54]^[55]^[56] As sensitive indicators of ecosystem health, mysids respond acutely to pollution and hypoxia, exhibiting behavioral disruptions, reduced growth, and elevated mortality at low dissolved oxygen levels (below 2 mg/L) and trace metal exposures, making them valuable for monitoring environmental degradation.^[57]^[58]^[59] Their population dynamics often follow boom-bust cycles, driven by resource availability and predation pressure, with rapid increases during favorable conditions followed by sharp declines that reflect broader ecological shifts.^[60]^[61]^[62]

Symbiotic and Parasitic Relationships

Mysids serve as important prey for a range of marine predators, including planktivorous fish such as Atlantic herring (Clupea harengus) and Pacific salmon (Oncorhynchus spp.), which consume them in significant proportions during certain life stages.^[63]^[64] Seabirds, exemplified by the marbled murrelet (Brachyramphus marmoratus), also incorporate mysids into their diets alongside other small crustaceans.^[65] Marine mammals, including humpback whales (Megaptera novaeangliae), have been documented consuming mysids, as evidenced by their presence in fecal samples.^[66] To evade these predators, gregarious mysids employ coordinated schooling behaviors, such as rapid increases in swimming speed or 90-degree escape trajectories relative to the attack direction, enhancing group survival.^[67]^[68] Within mysid populations, larger individuals act as predators on smaller conspecifics through cannibalism, which can regulate population densities and influence size structure.^[69] Symbiotic interactions involving mysids are predominantly commensal, such as associations with scyphozoan jellyfish (Cassiopea spp.), where mysids like Idiomysis gain transport and protection while inhabiting the jellyfish's tissues without apparent harm to the host.^[70]^[71] Mutualistic relationships are rarer but occur, for instance, when mysids excrete nutrients that enrich algal farms maintained by damselfish hosts, benefiting algal growth and indirectly supporting the fish.^[72] Mysids are susceptible to parasitic infestations that impact their fitness, including epibiotic protozoans such as ciliates, which colonize their exoskeletons and may increase drag or susceptibility to further infection.^[73] Trematode metacercariae, including progenetic forms, encyst in mysid tissues, potentially altering host behavior or energy allocation.^[74]^[75] Parasitic copepods, notably nicothoids, attach externally or internally, feeding on host embryos and larvae in the marsupium, thereby reducing fecundity by up to 36% in affected populations.^[76]^[77] These infestations collectively diminish reproduction and survival rates by impairing mobility, nutrient uptake, and immune responses. In aquaculture settings, mysids used as live feed can vector pathogens, such as mycobacteria, to cultured fish and invertebrates, potentially introducing infections into closed systems.^[78]

Taxonomy and Systematics

Higher Classification

Mysids belong to the phylum Arthropoda, subphylum Crustacea, class Malacostraca, subclass Eumalacostraca, and superorder Peracarida, where they are classified in the order Mysida.^[15] This placement reflects their shared peracarid traits, such as a marsupium for brooding young on the female's ventral side, and positions them alongside other orders like Amphipoda, Isopoda, and Cumacea within the diverse superorder Peracarida.^[15] The order Mysida comprises two families, Mysidae and Petalophthalmidae, encompassing a wide array of marine, freshwater, and subterranean species.^[15] Historically, mysids were grouped under the order Mysidacea, established by Pierre André Latreille in 1825 as part of the broader Schizopoda, which included euphausiaceans and leptostracans based on similarities in uropod and telson morphology.^[15]^[79] This classification persisted into the late 19th century, with refinements by Boas (1883) dividing Mysidacea into suborders Lophogastrida and Mysida, recognizing their distinct thoracic and abdominal features.^[79] Subsequent revisions, including Calman's (1904) introduction of Peracarida and later molecular analyses, led to the separation of Mysidacea into three distinct orders: Mysida, Lophogastrida, and Stygiomysida, with the latter comprising subterranean taxa sometimes combined with Mysida under a broader Mysidacea in older literature.^[15]^[79] Mysida are distinguished from related peracarid orders by their shrimp-like body form, featuring a carapace that covers the gills but does not fuse with the posterior thoracic segments, unlike the more rigid structures in some other malacostracans.^[15] In contrast, isopods exhibit dorso-ventral flattening and lack a distinct carapace, with their thoracic segments remaining largely free and no fusion akin to that seen in higher crustaceans.^[15] Amphipods, meanwhile, are laterally compressed, entirely lack a carapace, and are characterized by a hopping locomotion facilitated by their powerful pleopods, differing from the more swimming-oriented propulsion of mysids using their exopods and endopods.^[15] Molecular evidence, particularly from mitogenomic analyses of mitochondrial genomes across peracarid taxa, strongly supports the monophyly of Peracarida, including Mysida, by confirming shared derived characters in gene arrangement and phylogenetic clustering with high bootstrap support.^[80] These studies resolve earlier uncertainties from ribosomal DNA data and affirm Peracarida as a cohesive clade within Malacostraca, with Mysida positioned basally alongside Lophogastrida.^[80]^[15]

Diversity and Phylogeny

The order Mysida comprises two families: the speciose Mysidae, which includes the vast majority of genera and species, and the smaller, more specialized Petalophthalmidae, primarily adapted to deep-sea environments.^[15] Collectively, Mysida encompasses 183 genera and 1,233 species (as of November 2025), distributed across a wide array of habitats including marine, brackish, freshwater, and subterranean systems.^[81] High endemism is particularly notable in isolated habitats such as anchialine caves and ancient lake basins, where many species exhibit restricted ranges due to limited dispersal capabilities and habitat specificity.^[15] Phylogenetically, Mysida display distinct branches reflecting habitat transitions, with basal lineages often associated with freshwater invasions from marine ancestors, while derived radiations dominate modern marine ecosystems. Molecular and morphological analyses, particularly within genera like Mysis, reveal that continental and freshwater taxa form clades sister to circumarctic marine groups, underscoring multiple independent invasions into non-marine environments during evolutionary history. The order is positioned within the superorder Peracarida, with internal relationships supported by 18S rRNA sequences that highlight the monophyly of major subfamilies in Mysidae.^[82] The fossil record of Mysida is sparse but indicates an origin in the Jurassic, with the earliest known specimens, such as Elder unguiculata and Francocaris grimmi, preserved in Bavarian deposits.^[15] Modern diversity, particularly within key Mysidae subfamilies like Siriellinae, Gastrosaccinae, Mysinae, and Mysidellinae, emerged post-Cretaceous, as evidenced by the scarcity of pre-Cenozoic fossils and the radiation of statolith-bearing forms in Miocene Paratethys sediments.^[15]^[83] Recent taxonomic revisions, notably from 2015, have clarified evolutionary affinities by elevating the cave-adapted Stygiomysida—comprising 16 species in two families—as a distinct sister order to other peracarids, rather than a basal Mysida group, based on integrated molecular (18S rDNA, 28S rDNA) and morphological data.^[15] This separation emphasizes the specialized subterranean adaptations of Stygiomysida while affirming the broader phylogenetic context of Mysida within Peracarida.^[15]

Human Relevance

Ecological and Economic Uses

Mysids serve as important model organisms in ecotoxicology due to their sensitivity to pollutants, making them suitable for standardized toxicity assessments in marine and estuarine environments. The U.S. Environmental Protection Agency employs species such as Americamysis bahia in acute toxicity tests (OPPTS 850.1035) and chronic toxicity tests (OPPTS 850.1350), evaluating endpoints including survival, growth, and reproduction to determine chemical safety for regulatory purposes.^[84]^[85] These protocols highlight mysids' role in screening endocrine-disrupting chemicals and other contaminants at environmentally relevant concentrations.^[86]^[59] In aquaculture, mysids are valued as live feed for larval fish and shrimp, providing high nutritional content with elevated levels of proteins, lipids, and essential fatty acids that support rapid growth and survival. Mysid meal has been shown to replace up to 65.5% of fishmeal in diets for Pacific white shrimp (Penaeus vannamei) postlarvae, promoting comparable or enhanced performance.^[87] Compared to brine shrimp (Artemia), mysids offer superior nutrition for grouper larvae, improving feeding efficiency in hatchery systems.^[88] Their culture potential as alternative live feeds has been explored for sustainable aquaculture practices.^[44] Mysids function as effective bioindicators for monitoring water quality in coastal and estuarine habitats, owing to their responsiveness to pollutants, salinity variations, and turbidity. Zonal distributions of mysid species in estuaries reflect environmental gradients, enabling assessments of habitat health and contamination levels.^[89] Their use in such monitoring supports regulatory efforts to track anthropogenic impacts on aquatic ecosystems.^[90] In fisheries, mysids are harvested for use as bait, particularly in the North Atlantic region, where their swarms facilitate collection. On the Island of Jersey in the English Channel, mysids are processed into a bait paste called "cherve" sold to recreational and commercial fishermen.^[91] Extracts from mysids have shown potential biomedical applications, with studies examining their antimicrobial properties. Fermented mysid-based products, such as Thai kapi, demonstrate antibacterial activity against pathogens, attributed to bioactive peptides.^[92]

Conservation Concerns

Mysid populations in coastal and estuarine habitats face significant threats from habitat loss driven by human activities such as coastal development, urbanization, and pollution. These pressures degrade critical environments like seagrass meadows and mudflats, which serve as essential nurseries and feeding grounds, leading to reduced abundances in regions like the Mediterranean Sea where sediment loading and direct fluvial inputs exacerbate the issue.^[93] In continental systems, additional risks include hydrological alterations, deforestation of watersheds, and sediment intrusion, which disrupt benthic substrates and water quality, particularly affecting estuarine species.^[94] Climate change poses an escalating threat to mysids through warming waters that alter species distributions and physiological processes. In polar regions, declining sea ice—reduced by over 30% in the Arctic over the past four decades—endangers ice-associated species such as Mysis polaris and M. litoralis, which rely on sympagic habitats for survival and reproduction, potentially leading to range contractions or local extinctions. In 2025, Arctic sea ice reached a record low maximum extent of 14.33 million km², the lowest in the 47-year satellite record, further exacerbating risks to these sympagic species.^[95]^[96] Warmer temperatures also influence estuarine communities globally, with elevated salinities and heat stress impairing mitochondrial function and development in species like Neomysis integer, shifting community structures and favoring thermotolerant invaders.^[97] These changes highlight mysids' sensitivity as early indicators of climatic shifts in marine ecosystems.^[98] Invasive non-native mysids, such as Hemimysis anomala in the Laurentian Great Lakes, introduce competition and alter trophic dynamics, outcompeting native species like Mysis diluviana for resources and potentially reducing biodiversity in freshwater systems.^[53] This invasion, first detected in 2006, has spread across all five lakes, exacerbating ecosystem disruptions under concurrent stressors like climate variability. Overharvesting, primarily through incidental capture in shrimp trawls and targeted collection for aquaculture feed, impacts swarm-forming coastal species, with unregulated harvesting in some regions contributing to localized declines despite efforts toward sustainable practices.^[99] In continental contexts, illegal commerce further threatens rare taxa.^[94] Significant data gaps hinder comprehensive conservation, with IUCN Red List assessments available for fewer than 10% of the over 1,100 known mysid species worldwide, and most continental taxa (out of approximately 86 described) classified as Data Deficient due to limited ecological studies.^[94] Subterranean and anchialine species remain particularly unstudied, complicating threat evaluations and mitigation strategies, while only a handful—such as Diamysis pusilla, preliminarily assessed as Critically Endangered—have formal threat statuses in literature.^[94] Addressing these gaps requires expanded surveys and monitoring to inform targeted protections.

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[65]
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[66]
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[74]
(TREMATODA) FROM CANTERBURY - University of Canterbury
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[75]
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[76]
[PDF] MYSIDACEA) IN RELATION TO - NWFSC SDM Applications
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https://www.marinespecies.org/aphia.php?p=taxdetails&id=149668
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[84]
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[87]
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[90]
Mysida (Mysids) | Encyclopedia.com
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[91]
Characterization and in vitro biological activities of Thai traditional ...
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[92]
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[93]
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[94]
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[95]
[PDF] Climate Change and its Effect on Mitochondrial Function Among the ...
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Mysids have recently been used in fish farming projects as a live food resource, especially in freshwater fisheries.Missing: harvesting | Show results with:harvesting