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Amphipoda

Amphipoda is an order of malacostracan crustaceans within the superorder , distinguished by their laterally compressed bodies, absence of a , and shrimp-like form with leg-like appendages on most body segments. These small to moderately sized typically range from 1 mm to over 300 mm in length, though most species are under 20 mm, and exhibit swift, hopping locomotion adapted to diverse environments. Comprising over 10,800 described across more than 1,600 genera and 221 families as of 2025, Amphipoda represents one of the most diverse groups of crustaceans, with ongoing discoveries particularly in habitats. While predominantly , amphipods occupy a wide array of ecological niches, including freshwater , lakes, and moist terrestrial soils, where they require high humidity to prevent . In aquatic settings, they are often found among , under , or on the surface, serving as detritivores, herbivores, , or predators that play crucial roles in and as prey for , , and other . Amphipods are notable for their brood-carrying behavior, where females retain embryos in a marsupium formed by oostegites on their thoracic legs, leading to direct development without free larval stages in many . This reproductive strategy contributes to their rapid population growth and adaptability, with some producing multiple generations per year in favorable conditions. Ecologically, they influence benthic community structure through bioturbation and grazing, and certain , like gammarids, are key indicators of in programs due to their sensitivity to .

Taxonomy and Classification

Taxonomic History

The taxonomic history of Amphipoda traces back to Carl Linnaeus, who in 1758 included several amphipod species, such as Gammarus pulex (originally Cancer pulex), within the broad genus Cancer in Systema Naturae, reflecting the limited understanding of crustacean diversity at the time. This placement grouped amphipods with crabs and other decapods, overlooking their distinct laterally compressed body form and ambulatory pereopods. In the early 19th century, advanced the classification by coining the term "Amphipoda" in 1814, establishing it as a family-level group in his article on crustaceology for Brewster's Edinburgh Encyclopaedia, based on observations of their equal-sized thoracic segments and hopping locomotion. Leach's work separated amphipods from other peracarids and decapods, emphasizing morphological traits like the absence of a and the structure of their appendages. Subsequently, Alphonse Milne-Edwards in 1840 elevated Amphipoda to full order status within the class in his comprehensive Histoire Naturelle des Crustacés, integrating new collections from global expeditions and solidifying its position alongside orders like and . The 20th century brought major revisions through regional monographs and expedition reports. K.H. Barnard in 1916 published a seminal study on the amphipod of , describing numerous species and proposing refinements to subordinal divisions within Gammaridea, which highlighted regional and influenced global keys for identification. Similarly, K. Stephensen in 1923 contributed to suborder classifications in his report on the Danish Ingolf Expedition, detailing Hyperiidea and Gammaridea structures and establishing several families, such as Brachyscelidae, based on pelagic and benthic collections from the North Atlantic. Discoveries of deep-sea and parasitic amphipods, particularly from expeditions like the HMS Challenger (1872–1876) and later deep-water surveys, prompted significant taxonomic shifts by revealing highly specialized forms, such as and bopyrid-associated parasites, which necessitated new families and genera to accommodate their aberrant morphologies. These findings expanded the perceived ecological and morphological diversity, leading to the recognition of over 10,000 described species by the , as cataloged in ongoing databases. A notable in the 1980s centered on the status of Caprellidea relative to Gammaridea, driven by morphological analyses showing distinct body plans and appendage reductions in caprellids (e.g., fused segments and reduced coxae), though later revisions in subordinated Caprellidea to superfamily rank within Corophiidea (now part of Senticaudata).

Current Classification and Diversity

The order Amphipoda is classified within the superorder of the class in the phylum Arthropoda. Modern recognizes six suborders, based on the 2017 phylogenetic revision by Lowry and : Amphilochidea, Colomastigidea, Hyperiidea, Hyperiopsidea, Pseudingolfiellidea, and Senticaudata. This classification separated the former suborder Ingolfiellidea as the new order Ingolfiellida. The largest group, Senticaudata (encompassing much of the former Gammaridea and Corophiidea), includes predominantly benthic and accounts for approximately 6,100 described , primarily in marine and freshwater environments; it includes the superfamily Caprelloidea (formerly Caprellidea, known as skeleton shrimps for their slender, plant-like bodies), with roughly 1,200 often associated with epibiotic lifestyles on algae and animals. Amphilochidea comprises around 4,300 across diverse ecologies. Hyperiidea includes approximately 300 pelagic adapted to open ocean habitats. Colomastigidea features about 60 , Hyperiopsidea about 15 , and Pseudingolfiellidea only 4 , confined to and habitats. Prominent families exemplify this diversity: Gammaridae within Senticaudata, featuring genera like that are widespread in freshwater and coastal systems; Hyperiidae in Hyperiidea, with genera such as Hyperia known for symbiotic associations with ; and Caprellidae in Caprelloidea (Senticaudata), including genera like Caprella that dominate fouling communities. These families highlight the order's ecological breadth, from detritivores to predators and commensals. As of 2025, approximately 10,800 of Amphipoda have been described, with an estimated 2,000 additional undescribed based on ongoing surveys and molecular inventories; the greatest occurs in coastal habitats, where over 70% of are recorded. Recent taxonomic revisions, particularly from molecular phylogenies in the , have restructured the group by splitting the traditional Gammaridea into Senticaudata and Amphilochidea, elevating several families (e.g., within gammaroideans) through DNA-based evidence of cryptic and convergent morphologies.

Morphology and Anatomy

External Features

Amphipods exhibit a distinctive laterally compressed , which contrasts with the dorso-ventrally flattened form typical of many other peracarid crustaceans such as isopods. The body is divided into three main tagmata: the head, or cephalon; the , known as the pereon with seven segments; and the abdomen, comprising the pleon with six segments and a terminal urosome. This segmentation supports diverse locomotor and sensory functions, with the pereon bearing seven pairs of thoracic appendages and the pleon housing specialized abdominal limbs. The absence of a leaves the body segments exposed, allowing for flexibility in movement. The head features two pairs of antennae that play key roles in sensory perception. Antenna 1 is typically longer than antenna 2 and uniramous, aiding in mechanoreception and navigation. Antenna 2 is shorter and equipped with dense arrays of chemosensory setae, primarily for detecting chemical cues in the . The first two pairs of pereopods, termed gnathopods, are modified into subchelate grasping structures, with the propodus and dactylus forming a pincer-like apparatus for feeding and manipulation. Pereopods 3 through 7 are ambulatory, adapted for walking, clinging to substrates, or burrowing in benthic species. Abdominal appendages further diversify function across habitats. The pleon bears three pairs of biramous pleopods, which in free-swimming forms like hyperiids facilitate through undulating motions, while in benthic taxa they often contribute to by generating water currents. The urosome includes three pairs of uropods and a , forming a fan-like used for steering, braking, or leaping in terrestrial and semi-terrestrial species. Eyes in amphipods are and typically sessile, embedded directly on the cephalon for panoramic in gammaridean forms. In pelagic hyperiids, eyes may be enlarged and bi-lobed, with dorsal portions adapted for detecting light, though remaining sessile rather than on stalks. The is often translucent, minimizing visibility in aquatic environments, or bears pigments for disruptive against varied backgrounds. Sexual dimorphism is prominent in appendage morphology. Males possess enlarged, robust gnathopods, particularly the second pair, which are used in mate grasping and combat. Females develop oostegites, plate-like extensions from the coxae of pereopods 2–5, which form a brood pouch for carrying developing embryos. These differences enhance reproductive roles without altering the overall body proportions significantly.

Internal Systems

The digestive system of amphipods is a straight, differentiated tube comprising a , , and , adapted for processing diverse food sources ranging from to live prey. The , lined with , includes an leading to a cardiac equipped with a gastric mill—a set of and crushing plates that grind ingested material, with variations in structure correlating to feeding habits such as scavenging or predation. The , or mesenteron, serves as the primary site for absorption, while the facilitates water reabsorption and waste expulsion. The , a paired digestive gland attached to the , produces enzymes for breakdown and aids in and . Amphipods possess an open circulatory system characterized by a dorsal heart that pumps hemolymph through arteries into body sinuses. The heart features three pairs of incurrent ostia for hemolymph entry and typically gives rise to multiple cardiac arteries, with the number varying by species and directing flow to gills on pereopods 2 through 6 (or 7 in some species) for oxygenation. Hemolymph, containing hemocyanin as the oxygen carrier, bathes tissues directly before returning to the heart via open sinuses. Respiration in amphipods occurs primarily through branchial , which facilitate oxygen uptake and are to their aquatic and semi-terrestrial lifestyles. In gammaridean amphipods, these consist of simple sac-like or plate-like epipods attached to the coxae of pereopods 2 to 6, sometimes supplemented by a small on the coxa or sternal of segment 7 in certain species. The also support ionoregulation, with water currents generated by limb movements enhancing . In some terrestrial talitrid amphipods, modified function as lung-like structures for air , retaining moisture through vascularization. The of amphipods features a , or , located dorsally and bent anteriorly, which integrates sensory input and coordinates . This connects via circumesophageal connectives to a subesophageal ganglion and extends posteriorly as a ventral nerve chain with segmental ganglia innervating appendages and viscera. Statocysts, paired sensory organs in the first antennae (antennules), provide balance and orientation cues, particularly in pelagic or swimming species. Excretion and in amphipods are managed by paired antennal glands, located in the head and opening near the antennal bases. These glands filter , reabsorb ions, and produce to maintain internal , enabling adaptation to hypo- and hypersaline environments from marine to freshwater habitats. In species like Corophium volutator, the glands exhibit fine-structural features such as podocytes for , supporting efficient waste removal and ionic balance.

Size, Growth, and Variation

Body Size Ranges

Amphipods display considerable variation in body size, with most species ranging from 1 to 30 mm in body length, though extremes include forms as small as less than 1 mm and deep-sea giants reaching up to 340 mm. The smallest species, such as certain taxa in the family Melitidae, inhabit sandy sediments and rarely exceed 1.5 mm, adapting to confined spaces through miniaturization. In contrast, the largest extant amphipod, Alicella gigantea, a deep-sea , attains lengths of about 34 cm, exemplifying abyssal gigantism in the order; recent studies as of 2025 suggest this species may inhabit over half of the world's deep seafloor. Size differences are evident across suborders. Senticaudata (formerly classified under Gammaridea), the most diverse group, typically measure 2 to 20 mm, encompassing both marine and freshwater forms like the common gammarids. Hyperiidea, primarily pelagic, range from 1-2 mm in smaller species to over 20 mm, with some reaching up to over 100 mm, such as Cystisoma spp., in oceanic environments. Caprellidea, known as skeleton shrimps, are slender and elongated, generally 5 to 40 mm long, though their thin build gives them a delicate appearance despite comparable lengths to senticaudatans. Growth in amphipods occurs through episodic molting (ecdysis), with juveniles typically undergoing ecdysis every 1 to 2 weeks under optimal conditions, allowing rapid size increases of 20-50% per cycle. This frequency decreases with age and body size, extending to monthly intervals in adults. Environmental factors strongly influence growth patterns: higher temperatures accelerate molting rates, as seen in Gammarus species where intervals shorten from weeks at 8°C to days at 20°C; optimal salinities (around 10-35 ppt) enhance survival and growth in euryhaline taxa like Hyale crassicomis, while extremes reduce ecdysis success; and nutritional quality, particularly protein-rich diets, supports faster development and larger final sizes in juveniles. Larger body sizes often correlate with scaled anatomical features, such as extended appendages for locomotion in deep-sea species.

Morphological Adaptations

Amphipods exhibit a remarkable array of morphological adaptations that enable them to exploit diverse habitats, from benthic sediments to open ocean waters and terrestrial environments. These variations often involve modifications to appendages, body shape, sensory structures, and , reflecting selective pressures from type, predation, and resource availability. For instance, benthic species have evolved specialized pereopods for burrowing or tube , while pelagic forms prioritize streamlined profiles and enhanced for in the . In benthic habitats, sand-dwelling amphipods such as those in the family Haustoriidae possess robust, elongate posterior pereopods adapted for burrowing through loose . These appendages feature strong, curved dactyli and dense setae that facilitate propulsion and displacement, allowing efficient subsurface and refuge from surface predators. Similarly, corophiid amphipods, including in Corophium, utilize specialized setae on their anterior pereopods and gnathopods to gather and bind particulate materials with silk-like secretions from cephalic glands, constructing protective U-shaped in soft substrates. These , often incorporating grains or , provide stability and capabilities for deposit feeding. Pelagic hyperiid amphipods display streamlined, bodies that reduce drag during active swimming and diel vertical migrations, spanning depths from surface waters to over 1000 meters. This body plan, combined with reduced segmentation in the pleon and abbreviated tail fan, enhances hydrodynamic efficiency for pursuing gelatinous prey. Prominent adaptations include massively enlarged compound eyes, which in species like Hyperia galba occupy much of the head and feature multifaceted ommatidia optimized for detecting bioluminescent signals in low-light midwater environments, aiding in host location and predator avoidance. Parasitic hyperiids, particularly those associating with hosts, undergo significant morphological reductions to facilitate attachment and nutrient uptake. Juveniles and adults often exhibit abbreviated body segmentation, with fused pleonal somites and diminutive uropods, minimizing exposure while embedded in host tissues. Specialized hooks on gnathopods and pereopods, as seen in genera like Hyperia, enable secure anchorage to medusae, allowing parasitoids to feed on host fluids without triggering escape responses. These modifications represent a derived state within Hyperiidea, contrasting with free-living relatives. Terrestrial talitrid amphipods, such as beach fleas in the family , have developed a thickened to minimize loss in humid but desiccating soils and leaf litter. This integumentary reinforcement, coupled with a trend toward reduced surface area compared to aquatic kin, supports via branchial and in air. For rapid escape from predators, they possess enlarged, muscular posterior pereopods (pereopods 6 and 7) with elongated propodi and dactyli, enabling saltatory jumps up to several body lengths. In symbiotic associations, caprellid amphipods on algal hosts exhibit slender, elongate bodies with sparse segmentation and fine, hair-like setae that mimic the filamentous structure of macroalgae like , providing effective against visual predators. This morphological , observed in species such as Caprella dilatata, allows them to blend seamlessly with host surfaces, reducing detection while grazing on epiphytes or benefiting from host-derived shelter.

Reproduction and Life Cycle

Reproductive Strategies

Amphipods are predominantly dioecious, with distinct male and female es that exhibit , such as differences in gnathopods or body proportions; body size may be larger in either sex depending on the . Males employ precopulatory guarding as a primary strategy, grasping receptive females with their enlarged gnathopods for periods ranging from hours to days prior to the female's molt, ensuring priority access to fertilization. This behavior is widespread across gammaridean amphipods and aligns with the brief window of female receptivity immediately following . Fertilization in amphipods is internal, typically involving the transfer of spermatophores from the male to the female's genital openings or marsupial region during or shortly after mate guarding. In many species, such as those in the Gammaridae, males deposit paired spermatophores externally on the female's ventral surface, from which sperm migrate to fertilize eggs released into the marsupium. Following fertilization, females brood the embryos in a ventral marsupium formed by oostegites, providing protection and oxygenation for 2–8 weeks depending on and species, with shorter durations (e.g., 10–12 days at 25°C in Parhyale hawaiensis) in warmer conditions. Asexual reproduction is rare in Amphipoda but documented as in select , such as the groundwater-dwelling Stygobromus hayi, where unfertilized eggs develop into females, potentially aiding persistence in isolated habitats. varies widely, with brood sizes typically ranging from 5 to 200 eggs per female, influenced by body size, environmental factors like temperature, and habitat; for instance, smaller freshwater like Hyalella longistila average around 13 eggs per brood. In temperate to warm waters, females often produce multiple broods annually (up to 4–7 per lifetime in like Grandidierella japonica), enabling rapid population growth.

Developmental Stages

Amphipods exhibit direct development, lacking a free-living larval , with embryos developing within the female's marsupium until hatching as miniature adults known as juveniles. These juveniles possess all major appendages and body segments typical of adults, though at a reduced size, and emerge from the marsupium fully formed for independent life. The process begins with lecithotrophic eggs, nourished solely by yolk reserves during intra-marsupial embryogenesis, which typically lasts 10–12 days at 25°C in model species like Parhyale hawaiensis. Post-hatching, juveniles undergo sequential molts to reach maturity, with 5–7 instars common in many gammaridean species, though up to 15 molts occur in some depending on environmental conditions and species. Time to maturity varies from 6 weeks at 18–25°C in tropical or subtropical species to 3 months or longer in temperate or cold-water forms, with higher temperatures accelerating molt frequency and growth rates. Each molt adds size and refines structures, but the overall pattern remains anamorphic, with no major metamorphic shifts in non-parasitic taxa. Environmental factors significantly influence ; optimal (e.g., 20–35‰ in marine ) maximizes success, while deviations reduce viability and survival rates in the marsupium. Temperature modulates developmental pace, with rates doubling from 10°C to 20°C in like Echinogammarus marinus, though extremes can cause developmental arrest or mortality. Variations exist among suborders; hyperiid amphipods often release juveniles into brief planktonic "physosoma" stages before adopting a pelagic lifestyle, differing from the benthic direct release in gammarids. Parasitic amphipods, such as Hyperia galba, undergo during host association, with juveniles transforming morphologically after attachment to medusae or other hosts.

Habitats and Distribution

Marine Environments

Amphipods are predominantly organisms, with approximately 80% of the over 10,000 described inhabiting environments, spanning from coastal zones to the deepest trenches. This dominance underscores their role as key components of ecosystems, where they exhibit remarkable adaptability to diverse conditions including varying salinities, pressures, and temperatures. In intertidal zones, amphipods such as those in the genus Orchestia (family Talitridae) are common burrowing forms, excavating tunnels in sand or wrack to evade and predators during low . These semi-terrestrial thrive in the dynamic littoral , contributing to nutrient cycling through detritivory. Pelagic amphipods, particularly the hyperiid suborder, inhabit open ocean waters and often associate with gelatinous like and salps, using them as hosts or refuges. Many hyperiids undertake diel vertical migrations, descending to deeper waters during the day and ascending at night to follow prey and avoid visual predators, a that facilitates their widespread . Benthic amphipods occupy seafloor habitats, with epifaunal species clinging to and seagrasses for shelter and , while infaunal forms burrow into sediments to feed on . In deep-sea environments, such as abyssal plains and hadal trenches, exceptionally large "giant" or "supergiant" amphipods (e.g., Alicella gigantea) reach lengths up to 34 cm, scavenging organic falls like carcasses; recent studies as of 2025 indicate that A. gigantea may inhabit up to 59% of the deep-sea floor, suggesting it is more widespread than previously thought. Amphipods exhibit distinct zonation patterns across marine depths: the hosts high species diversity due to heterogeneous substrates, the neritic shelf supports abundant epibenthic communities, and the bathyal slope features specialized deep-water forms adapted to diminishing light and oxygen. In polar regions, species like Themisto (Hyperiidae) dominate pelagic assemblages in the and , enduring extreme cold and seasonal ice cover. Key adaptations enable amphipods to exploit marine salinity gradients, particularly in coastal and estuarine interfaces, through efficient osmoregulation via active ion transport in the gills and gut. Bioluminescence, though rare, occurs in some abyssal species (e.g., certain hyperiids), serving as a defense mechanism against predators in the dark deep sea.

Freshwater and Terrestrial Habitats

Approximately 20% of all amphipod , or about 1,900-2,000 taxa as of 2025, inhabit freshwater environments worldwide, representing a significant departure from the predominantly nature of the . These thrive primarily in cool, temperate regions, with roughly 70% of the global freshwater diversity concentrated in the Palaearctic realm, including rivers, lakes, and subterranean habitats. Diversity is particularly elevated in running waters and systems, where amphipods often serve as key components of benthic communities. Surface freshwater ecosystems are dominated by gammarid amphipods, especially from the genus , which are widespread in lotic and lentic habitats across the ; for instance, Gammarus pulex is a prevalent species in European rivers and streams, exhibiting broad tolerance to varying flow regimes and temperatures. In contrast, and hyporheic zones—transitional areas between surface and subsurface waters—host specialized taxa like those in the genus Niphargus (family Niphargidae), the most species-rich freshwater amphipod group with over 450 described species as of 2025, many of which demonstrate physiological adaptations to low-oxygen conditions through enhanced respiratory efficiency and behavioral microhabitat selection. These subterranean forms often show reduced pigmentation and elongated appendages suited to confined, dark environments. Invasive Ponto-Caspian amphipods have notably altered freshwater distributions in recent decades, with species such as Dikerogammarus villosus spreading rapidly across European waterways primarily through ship ballast water and recreational boating, where their predatory behavior has led to the local elimination of native gammarids like Gammarus duebeni. This invasion highlights the vulnerability of temperate freshwater systems to non-native introductions, often resulting in shifts toward more aggressive, less diverse amphipod assemblages. Terrestrial habitats support a much smaller proportion of amphipod diversity, limited almost exclusively to the family , which includes "landhoppers" adapted to life above water in moist, vegetated settings such as forest leaf litter, coastal dunes, and supralittoral zones. These environments are predominantly coastal or humid inland forests, with the highest concentrations in the and , where talitrids exploit detrital resources in stable, shaded microhabitats. Unlike their aquatic relatives, terrestrial talitrids feature morphological adaptations including reduced or modified gills with expanded surface areas for cutaneous air breathing, coupled with behavioral strategies like burrowing and nocturnal to minimize and maintain branchial hydration. Globally, freshwater amphipod richness aligns closely with temperate climatic zones, reflecting preferences for cooler waters and stable , whereas terrestrial forms remain largely confined to coastal and peri-humid ecosystems, underscoring the order's limited success in fully conquering inland terrestrial niches.

Ecology and Behavior

Feeding and Foraging

Amphipods predominantly function as detritivores, shredding and consuming such as decaying material, fragments, and microbial films in and freshwater sediments using their powerful mandibles and maxillipeds. This feeding mode is especially prevalent among benthic gammaridean amphipods, which into soft substrates to access and process , facilitating nutrient recycling in ecosystems. Scavenging behaviors complement detritivory, with like lysianassoids rapidly aggregating on carrion to consume larger remains, often detected through baited traps in deep-sea studies. Herbivory occurs in several groups, particularly caprellid amphipods, which graze on epiphytic and biofilms coating substrates like seagrasses and hydroids by scraping with specialized gnathopods and pereopods. For instance, Caprella penantis combines scraping with occasional filter-feeding to harvest suspended , adapting to phytal habitats where plant-based resources abound. Carnivorous and omnivorous strategies are evident in predatory species that ambush or pursue small , including copepods, polychaetes, and juvenile crustaceans, using appendages to grasp and tear prey. Planktonic hyperiids, such as Themisto species, exemplify omnivory by preying on like euphausiids and copepods, with gut analyses revealing a broad spectrum of animal matter. Foraging in amphipods often involves nocturnal activity to reduce predation risk, with individuals emerging from refuges at dusk to actively search sediments or water columns for food. Planktonic forms engage in swarming behaviors that enhance encounter rates with patchy resources or mates, while some caprellids adopt symbiotic associations on host organisms, feeding on associated fouling communities or host exudates without harming the host. Gut content examinations, supplemented by stable isotope and DNA metabarcoding analyses, consistently demonstrate mixed diets across species, underscoring their opportunistic nature and primary role as basal consumers in trophic webs, though some shift to higher levels when animal prey is available.

Interactions and Roles in Ecosystems

Amphipods serve as a critical prey base in various ecosystems, supporting higher trophic levels such as , , and marine mammals. In polar regions, sympagic amphipods like Apherusa glacialis play a krill-like role, with an estimated 60 million tonnes consumed annually by , , and mammals, contributing substantially to their diets. In benthic marine environments, amphipods form an important link between primary producers and secondary consumers, serving as prey for , seabirds, and cetaceans, including gray whales that target dense amphipod patches. In freshwater streams, amphipods can constitute a significant portion of macroinvertebrate biomass, often exceeding 25% in some systems, making them a key food resource for predatory and invertebrates. Certain amphipod species act as predators within food webs, preying on smaller zooplankton like copepods, which enhances their role in regulating lower trophic levels. For instance, the invasive amphipod Themisto libellula primarily consumes copepodite stages of Calanus finmarchicus in marine systems. Hyperiid amphipods often exhibit parasitic or commensal associations with gelatinous zooplankton, such as medusae, where they inhabit and feed on host tissues, sometimes acting as a trophic link to fish predators. Amphipods themselves serve as intermediate hosts for diverse parasites, including trematodes and cestodes, which can influence population dynamics and trophic transmission in aquatic ecosystems. As decomposers, amphipods facilitate nutrient cycling by processing , particularly in freshwater and coastal systems where species like gammarids break down leaf and . Gammarid amphipods, such as Gammarus fossarum, play a central role in decomposition, contributing to energy transfer and nutrient release in streams. They are widely used as bioindicators of due to their sensitivity to contaminants; for example, Gammarus species are employed in toxicity tests to assess metal and chemical impacts in aquatic environments. Amphipods engage in various symbiotic interactions that shape community structure. Leucothoid amphipods live commensally within sponges, filtering particles from host-generated water currents without apparent harm to the sponge. Some species form mutualistic relationships with macroalgae, where amphipods graze on epiphytes, indirectly benefiting algal growth and health. In community dynamics, amphipods often function as keystone species, particularly in leaf litter breakdown processes that sustain detrital food webs in lotic ecosystems. However, invasive amphipods can disrupt biodiversity; for example, Echinogammarus ischnus has displaced native Gammarus fasciatus in the Great Lakes, altering benthic community composition and potentially reducing native species diversity. Such invasions may also impact ecosystem functions like detritus processing and prey availability for native predators.

Evolutionary History

Fossil Record

The fossil record of Amphipoda is exceedingly sparse, owing to the peracaridans' predominantly soft-bodied construction, which rarely preserves well outside exceptional Lagerstätten; over 35 species have been described as of 2025, far fewer than their estimated 10,000+ extant species. Fossils are typically preserved as compressions in fine-grained sedimentary deposits or as three-dimensional inclusions in , with the latter providing the most detailed morphological insights into appendages and body segmentation. This limited record contrasts sharply with the group's inferred ancient origins, highlighting significant gaps, particularly before the , where no confirmed amphipod fossils exist despite phylogenetic estimates placing their emergence in the , potentially as early as the around 320 million years ago. The earliest undisputed amphipod fossils appear in the , marking the onset of their documented diversification. The oldest known species, Gammaroidorum vonki, comes from Lower marine deposits in the Wealden Group of southeast , dating to approximately 130 million years ago; this diminutive gammaridean form, about 3 mm long, suggests early adaptation to shallow coastal environments. occurrences remain rare overall, with no verified amber inclusions from the , though the record hints at emerging ecological roles in marine ecosystems, including possible associations with gammaridean lineages that dominate modern diversity. Notably, amphipods show no evidence of being uniquely impacted by mass extinction events during this period, likely due to their benthic and opportunistic habits allowing persistence through environmental upheavals. The era reveals a marked increase in fossil abundance and variety, underscoring post-Mesozoic radiation into diverse habitats. Eocene deposits, particularly the renowned (ca. 44–38 Ma), yield the richest assemblages, including over a dozen of terrestrial and freshwater forms such as Palaeogammarus and Synurella aliciae, often preserved in groups suggestive of precopulatory behaviors or mass drownings in resin-trapped wetlands. These inclusions document early colonization of non-marine environments by talitrid and gammarid relatives. records (ca. 23–5 Ma) further expand this pattern, with fossils from amber in revealing blind subterranean talitrids like Caecorchestia bousfieldi and from revealing Ponto-Caspian gammaroids, alongside marine compressions indicating adaptation to brackish and possibly deeper-water settings. In 2024, two new of fossil Ponto-Caspian gammaroids, Pontogammarus adamantis and Obesogammarus georgianus sp. nov., were described from (ca. 10.5 Ma) amber in eastern , providing further evidence of early diversification in non-marine environments. Overall, the post-Paleogene proliferation aligns with global climatic shifts that opened new ecological niches, though the record remains biased toward amber-preserved coastal and inland taxa.

Phylogenetic Relationships

Amphipoda belongs to the superorder within the subclass of the class , and is consistently recovered as monophyletic in both morphological and molecular analyses, often including or sister to the clade Mancoida (encompassing and related taxa). Within , Amphipoda is positioned as the to , supported by shared morphological features such as the ventral brood pouch (marsupium) derived from oostegites on the pereopods, which evolved once in the common ancestor of for protecting developing embryos. This placement is reinforced by the loss of the in Amphipoda, a derived trait distinguishing it from more basal peracarids like mysids that retain a partial covering the gills. Internally, the phylogeny of Amphipoda reveals Hyperiidea as a basal suborder, comprising pelagic taxa that diverged early from the lineage leading to benthic groups. The traditional suborder Gammaridea is paraphyletic, with various lineages nested among other groups, while Caprellidea (skeleton shrimps) represents a derived originating from within corophiid gammarideans through adaptations like body elongation and pereopod reduction for clinging to hosts. Molecular evidence from 18S rRNA and mitochondrial (mitogenome) studies in the and has driven revisions to amphipod subordinal classification, confirming the non-monophyly of Gammaridea and supporting the elevation of groups like Ingolfiellida to ordinal status. These analyses estimate the divergence of Amphipoda from close peracarid relatives around 400 million years ago during the , aligning with early peracarid fossils and marking the onset of their diversification. Key evolutionary traits in Amphipoda include the refinement of the brood pouch for direct development without free larval stages, inherited from peracarid ancestors but modified for oostegal protection in females. The complete loss of the , unlike in isopods where it forms a partial shield, facilitated greater body flexibility and is considered a synapomorphy for amphipod locomotion and habitat adaptation. Controversies persist regarding the exact position of Amphipoda relative to within , with some early molecular studies suggesting mysids as closer to eucarids, potentially rendering paraphyletic. However, recent phylogenomic analyses using hundreds of nuclear genes in 2023 have confirmed the monophyly of (including Amphipoda and ) and resolved Amphipoda as sister to a -Isopoda clade in most models, resolving prior conflicts through improved taxon sampling and site-heterogeneous models. A November 2025 phylogenomic , the most comprehensive to date, further supports monophyly and this sister-group relationship using extensive datasets.

Human Relevance

Economic and Scientific Uses

Amphipods, particularly species in the genus Gammarus, serve as important model organisms in due to their sensitivity to environmental contaminants and relatively short life cycles, which facilitate studies. For instance, G. pulex reaches within approximately 130 days under controlled conditions, enabling efficient assessment of chemical impacts on reproduction and survival. These amphipods are recommended for standardized toxicity tests, including guidelines for sediment-water assays using spiked sediments to evaluate persistent chemicals' effects on adult reproduction. Comprehensive reviews highlight over 200 studies using Gammarus spp. to measure endpoints like , , and sublethal responses in freshwater ecosystems. In aquaculture, amphipods are valued as a high-protein feed source for fish and shrimp farming, offering nutritional benefits such as elevated levels of polyunsaturated fatty acids. Cultured species like Parhyale hawaiensis provide essential omega-3 fatty acids, making them a viable alternative to traditional feeds in experimental diets for species including and . Similarly, Bemlos quadrimanus demonstrates potential to replace fishmeal in shrimp diets, with protein levels exceeding 50% when grown on low-cost substrates like macroalgae. These attributes support sustainable production, as amphipods can be mass-reared in simple systems, enhancing digestibility and growth rates in aquaculture operations. Biomedically, amphipods contribute to through enzyme extracts with specialized properties. For example, a psychrophilic aspartic isolated from the freshwater amphipod bakhteyaricus exhibits high activity at low temperatures (optimal at 20°C) and stability across a broad range, suitable for industrial applications in and detergents. Deep-sea species like Hirondellea gigas produce hydrolases that degrade complex carbohydrates, offering potential for production and due to their efficiency in extreme conditions. derived from amphipod exoskeletons has shown antifungal activity against pathogens like Fusarium oxysporum, supporting applications in crop protection. Scientifically, amphipods act as sentinels for ocean health, particularly in microplastic pollution since studies intensified around 2015. Benthic and pelagic species, including Gammarus setosus in regions and lysianassoid amphipods in deep-sea trenches, readily ingest , accumulating particles at higher levels than in surrounding environments. This enables assessment of transfer through food webs, with laboratory exposures revealing impacts on and leaf-shredding in species like G. duebeni. Their role as ecological indicators extends to broader , where high abundances signal enrichment or . A 2025 evidence review further underscores their sensitivity to contaminants like oil and metals in marine environments. Commercially, amphipods are harvested as bait for fishing, with freshwater Gammarus spp. commonly used in angling due to their natural abundance in streams. Skeleton shrimp (Caprella spp.) are popular in the ornamental trade for aquaria, valued for their active behavior and role in maintaining clean substrates by grazing diatoms. These uses leverage amphipods' availability and low cost, supporting both recreational and educational markets.

Impacts and Management

Amphipods exert significant ecological impacts through that disrupt native communities and ecosystem processes. For instance, the Ponto-Caspian amphipod , known as the killer shrimp, has invaded European freshwaters, where it preys on native and alters processing, with lower per capita shredding rates than natives but potentially greater total processing due to high abundance in affected streams. Recent 2025 studies highlight its variable dispersal potential driven by competition, complicating management. This invasion can lead to decreased and impaired nutrient cycling, with cascading effects on higher trophic levels such as populations. Similarly, other non-native amphipods like Gammarus tigrinus compete with indigenous species, contributing to shifts in community structure in the . In human-impacted environments, amphipods serve as sensitive bioindicators of , particularly for , , and emerging contaminants. Following the 1999 Erika off , amphipod assemblages in soft-sediment habitats showed marked declines in abundance and diversity compared to more resilient polychaetes, highlighting their utility for targeted post-spill monitoring. Studies have also demonstrated their effectiveness in detecting metal in marine sediments and perfluorochemicals in freshwater streams, where species like Gammarus pulex exhibit reduced growth and reproduction at low contaminant levels. In settings, amphipods often form communities on nets and structures, potentially increasing maintenance costs and harboring pathogens, though they also offer nutritional value as alternative protein sources for fish feed. Management of amphipod impacts focuses on preventing and controlling invasions while leveraging their indicator role for environmental protection. Strategies for invasive species like D. villosus include physical containment via barriers, reproductive removal through trapping, and emerging biotechnologies such as RNA interference (RNAi) and pheromone traps, though efficacy varies with site-specific factors like water flow and population density. Eradication efforts have had limited success due to the amphipods' high reproductive rates, prompting emphasis on early detection and public education to curb spread via boating and ballast water. For pollution monitoring, standardized protocols using amphipod community metrics are integrated into regulatory frameworks, such as those under the European Water Framework Directive, to assess habitat health and guide remediation. In aquaculture, integrated pest management combines mechanical cleaning with selective harvesting to mitigate fouling while exploring amphipods' potential in sustainable feed production.

References

  1. [1]
    Freshwater Amphipod Crustaceans (Gammaridae) of North America
    Amphipods belong to the order Amphipoda of the class Crustacea. Amphipoda is further classified in the superorder Peracarida of the subclass Malacostraca and is ...
  2. [2]
    general characteristics of crustaceans - File: <identifymed
    Order: Amphipoda. -- Primarily marine. They do not have a carapace. The eyes are sessile and the body is usually depressed. There is much variety exhibited ...Missing: physical | Show results with:physical
  3. [3]
    EENY-220/IN377: Terrestrial Amphipods or Lawn Shrimp (Crustacea
    Terrestrial amphipods, also called lawn shrimp, are shrimp-like crustacea that need moist habitats. They are elongate, lack a carapace, and have leg-like ...Missing: sources | Show results with:sources
  4. [4]
    Progress in the discovery of amphipod crustaceans - PMC
    Jul 11, 2018 · At present, amphipod crustaceans comprise 9,980 species, 1,664 genera, 444 subfamilies, and 221 families. Of these, 1,940 species (almost 20%) ...
  5. [5]
    [PDF] amphipod
    A large 28cm amphipod was found in the deep ocean. Most deep water amphipods are 2-3cm, but this one was 28-34cm.
  6. [6]
    Amphipoda - an overview | ScienceDirect Topics
    Amphipods are crustaceans, with about 7900 species, mainly marine, but also in freshwater and terrestrial habitats, known for swift locomotion.
  7. [7]
    Biota of Freshwater Ecosystems Identification Manual 5 - epa nepis
    In aquatic habitats in general, amphipods are usually found under gravels, dead leaves, grass, and other kinds of debris, or in masses of vegetation.
  8. [8]
    Diversity and Distribution of Freshwater Amphipod Species in ...
    Oct 29, 2014 · Amphipods are important for ecosystem processes and trophic dynamics in freshwater ecosystems and increasingly important for eco-monitoring and ...
  9. [9]
    Scud Scuds (Amphipoda) - Maine.gov
    Scuds are members of a bottom dwelling class of crustaceans called Malacostraca. This class also includes aquatic sow bugs (Isopoda), crayfish, and shrimps ( ...
  10. [10]
    [PDF] Pontogammarus robustoides (an amphipod, no common name)
    Four Ponto-Caspian and one American gammarid species (Crustacea, Amphipoda) recently invading Polish waters. Contributions to. Zoology 71(4):115-122. Page 17 ...
  11. [11]
  12. [12]
    History of Discovery of Parasitic Crustacea - PMC - PubMed Central
    The first significant contributor after Linnaeus was the English naturalist, William Elford Leach. Leach, who was one of the world's leading crustacean ...
  13. [13]
    World Amphipoda Database - WoRMS
    The order Amphipoda is very diverse and currently contains 10,819 species which are divided into six suborders: ... Accessed at https://www.marinespecies.org/ ...
  14. [14]
    (PDF) A Phylogeny and a New Classification of the Corophiidea ...
    Aug 7, 2025 · A Phylogeny and a New Classification of the Corophiidea Leach, 1814 (Amphipoda). January 2009; Journal of Crustacean Biology 23(May 2003):443 ...
  15. [15]
    World Register of Marine Species - Amphipoda - WoRMS
    Amphipoda · Suborder Amphilochidea · Suborder Colomastigidea · Suborder Hyperiidea · Suborder Hyperiopsidea · Suborder Senticaudata · Suborder Caprellidea accepted as ...
  16. [16]
    A Phylogeny and Classification of the Amphipoda ... - Magnolia Press
    May 11, 2017 · Abstract. A classification is proposed for the order Amphipoda. The Amphipoda includes six suborders, the Pseudingolfiellidea, Hyperiidea, ...
  17. [17]
    Marine amphipods as integral members of global ocean ecosystems
    The order Amphipoda consists of seven accepted suborders, six of which are found in marine environments: Amphilochidea, Colomastigidea, Hyperiidea, ...
  18. [18]
    (PDF) A Phylogeny and Classification of the Amphipoda with the ...
    Aug 7, 2025 · A classification is proposed for the order Amphipoda. The Amphipoda includes six suborders, the Pseudingolfiellidea, Hyperiidea, Colomastigidea, Hyperiopsidea, ...
  19. [19]
    Crangonyx - Lander University
    The mandible bears a movable tooth, the lacinia mobilis, between the molar and incisor. Amphipoda O. Amphipoda is the largest peracarid taxon with about 6000, ...
  20. [20]
    Amphipoda - Zooplankton - University of Tasmania, Australia
    Pereiopods 1 and 2 are typically modified as subchelate prehensile gnathopods (claw-bearing and grasping limbs) for processing food. Abdomen comprises 2 ...
  21. [21]
    A new species of Ampithoe (Amphipoda, Ampithoidae) from Korea ...
    The genus Ampithoe Leach, 1814 is a herbivorous amphipod group associated with algae and seagrasses in coastal regions worldwide (Myers and Lowry 2003; Shin et ...
  22. [22]
    Uropod - an overview | ScienceDirect Topics
    Amphipods are also unique in having three pairs of uropods (appendages at their tail end), which are used for pushing in aquatic species, and for jumping in ...
  23. [23]
    The “amphi”-brains of amphipods: new insights from the ...
    Posterolaterally attached to the lateral antenna 1 neuropil, the deutocerebral chemosensory lobe is innervated by a small lateral section of the antenna 1 nerve ...
  24. [24]
    A unique apposition compound eye in the mesopelagic hyperiid ...
    The hyperiid amphipod Paraphronima gracilis has a pair of bi-lobed apposition compound eyes, each with a large upward-looking portion and a small lateral ...
  25. [25]
    Hidden in Plain Sight: The Ecology and Physiology of Organismal ...
    For example, the chitinous cuticle of a transparent hyperiid amphipod (n = 1.55) reflects 0.5% of the light and would be visible at 625, 525, and 475 m for ...
  26. [26]
    The role of male posterior gnathopods revealed in Gammarus pulex
    Aug 7, 2025 · In most amphipods, the gnathopods (especially the second pair of gnathopods) show a pronounced sexual dimorphism and are generally larger, more ...Missing: oostegites | Show results with:oostegites
  27. [27]
    Variation in Gnathopod Morphology of Cave Amphipods and its use ...
    Jan 1, 2009 · Sex was based on primary sexual characteristics: a pair of penal papillae on the sternum of pleomere 7 for males and oöstegites (brood plates) ...Missing: oostegites | Show results with:oostegites
  28. [28]
    Digestive Anatomy of Hyalella azteca (Crustacea, Amphipoda)
    ABSTRACT. The digestive tract of the freshwater amphipod Hyalella azteca is a straight but differentiated tube consisting of foregut, midgut, and hindgut.Missing: system | Show results with:system
  29. [29]
    (PDF) Comparative fore-gut morphology of Antarctic Amphipoda ...
    Aug 6, 2025 · In amphipods, the morphology of the mouthparts and digestive system is known to vary according to lifestyle and feeding habits. Since ...
  30. [30]
    Digestive anatomy of Halella azteca (Crustacea, Amphipoda)
    The cuticular lining of the cardiac stomach is elaborated into a set of food-crushing plates and ossicles, the gastric mill, while the pyloric cuticle forms a ...
  31. [31]
    New digestive symbiosis in the hydrothermal vent amphipoda ...
    In amphipods, the midgut or mesenteron and the digestive gland are the major organs involved in digestion and absorption processes of nutrients. Their main ...
  32. [32]
    Functional anatomy of the midgut gland of Gammarus locusta ...
    The hepatopancreas is composed of two pairs of ... Functional anatomy of the midgut gland of Gammarus locusta (Crustacea: Amphipoda) ... The hepatopancreas ...
  33. [33]
    Comparative analysis of the circulatory system in Amphipoda ...
    Apr 6, 2007 · The heart is equipped with three pairs of incurrent ostia. The number of cardiac arteries that lead off the heart varies among species: in the ...
  34. [34]
    The structure and organization of the gills of gammaridean Amphipoda
    The gills of gammaridean amphipods consist of epipods on the coxa of peraeopods 2 to 6 and an exopod on the basis of peraeopod 7.Missing: system | Show results with:system
  35. [35]
    Gill function in the amphipod Megalorchestia (Orchestoidea ...
    Sep 26, 2025 · The gills of amphipods, like many crustaceans, are considered to be principal sites of ion and water exchange between the internal (i.e., ...
  36. [36]
    Anatomy of structures associated with air-breathing in Orchestia ...
    Chloride-ion-permeable areas were located using a silver-staining technique. All ten coxal gills appeared to be equally permeable to chloride ions after ...Missing: respiratory | Show results with:respiratory
  37. [37]
    New insights from the neuroanatomy of Parhyale hawaiensis (Dana ...
    Apr 18, 2019 · Results As in most amphipod crustaceans, the brain is dorsally bent out of the body axis with downward oriented lateral hemispheres of the ...
  38. [38]
    Microanatomy of the Central Nervous System of Gammarus setosus ...
    MICROANATOMY OF THE CENTRAL NERVOUS SYSTEM OF. GAMMARUS SETOSUS DEMENTIEVA (AMPHIPODA). THE SUPRAOESOPHAGEAL GANGLION (BRAIN). BY. B. R. MACPHERSON. Department ...
  39. [39]
    The general morphology and fine structure of the antennary gland of ...
    May 11, 2009 · The maxillary gland of Artemia salina is the only excretory system of a non-decapod which has been studied in detail (Tyson, 1968, 1969a, b).
  40. [40]
    Osmoregulation inCorophium curvispinum (Crustacea: Amphipoda ...
    Osmoregulation inCorophium curvispinum (Crustacea: Amphipoda), a recent ... antennal glands ofGammarus oceanicus (Segerstrale, 1947) Crustacea: Amphipoda.
  41. [41]
    The supergiant amphipod Alicella gigantea may inhabit over half of ...
    May 21, 2025 · This species has attracted widespread attention due to its significant gigantism with a maximum body length of 340 mm [8–10].
  42. [42]
    [PDF] CAPRELLIDEA - NMBAQC
    It is generally accepted that Caprellidea derived from Gammaridea (cf. Myers & Lowry, 2003; Ito et al., 2008), which are characterized by well developed ...Missing: separation | Show results with:separation
  43. [43]
    Effects of temperature, age and body size on moulting and growth in ...
    Maturation times increased from 96 days at 20.2°C to 355 days at 7.9°C in G. fossarum, and 85 days to 403 days in G. roeseli. 4. Over the range 3.8 ...
  44. [44]
    (PDF) Effects of temperature and salinity on survival and growth, of ...
    Aug 6, 2025 · At 25°C, the amphipods tolerated salinities between 10 and 50 (with >80% survival in 96 h). The 96‐h LS50 values at high and low salinity ranges ...
  45. [45]
    Growth, Molting Frequency, Heart Beat, Number of Eggs, and ...
    Effects of temperature, age and body size on moulting and growth in the freshwater amphipods Gammarus fossarum and G. roeseli · Environmental Science, Biology.
  46. [46]
    Phylogeny and a New Classification of the Corophiidea Leach, 1814 ...
    Leach (1814: 405) erected the Corophiidae (as the family Corophiini) where it has page placement priority over the family Caprellini. This error has since been ...
  47. [47]
    Functional differences between the extraordinary eyes of deep-sea ...
    May 29, 2024 · Hyperiid amphipods (Crustacea, Malacostraca, Peracarida) provide an ideal opportunity to investigate visual adaptations to the midwater.
  48. [48]
    The habits and feeding mechanism of the Amphipod Haustorius ...
    Aug 9, 2025 · These species are sand-dwelling filter and deposit feeders, but differ slightly in feeding strategy making them functionally distinct.
  49. [49]
    [PDF] Studies on the Biology of the Tube-Building Amphipod ... - DTIC
    In contrast, the morphology of newly hatched amphipods closely resembles the adults', except for their sexual characteristics, which develop gradually ...
  50. [50]
    Tube building by the amphipods Lembos websteri Bate and ...
    Corophium build tubes with openings raised above the substratum more commonly than Lembos, and will construct mats of tubes away from solid surfaces; Lembos ...Missing: setae | Show results with:setae
  51. [51]
    Phylogenetic analysis of lineage relationships among hyperiid ...
    Oct 30, 2007 · The pelagic amphipods, hyperiids, have highly divergent life histories and morphological attributes in comparison to more familiar benthic, ...
  52. [52]
    [PDF] Amphipod Parasites of Fishes of Canada
    STEPHENSEN, K. 1923. Amphipoda I, Crusta- cean, Malacostraca V. Danish Ingolf. Exped. 3 (8). 1944. Amphipoda ...
  53. [53]
    A method for eradicating amphipod parasites (hyperiidae) from host ...
    The use of diflubenzuron to eradicate hyperiid parasites from scyphomedusae is a safe and useful option when properly applied in a controlled environment.Missing: segmentation hooks
  54. [54]
    BIOLOGY OF TERRESTRIAL AMPHIPODS - Annual Reviews
    The enlargement of the gills, despite the apparent abundance of oxygen on land, may be due to an increase in the thickness of their cuticle to prevent ...
  55. [55]
    Terrestrial crustaceans (Arthropoda, Crustacea) - ZooKeys
    Jul 13, 2023 · The cuticle of the gill lamellae of almost all air-breathing terrestrial decapods is usually much thicker than that of their aquatic relatives ( ...
  56. [56]
    [PDF] Associations between amphipods (Crustacea: Amphipoda) and sea ...
    The low fecundity of the symbiotic amphipods is probably made possible by the protection afforded by the host. Moreover in O. recondita the reproductive ...
  57. [57]
    Population biology of sympatric species of Caprella (Amphipoda
    Significant temporal variations were recorded for the three species with higher densities in spring and summer that are related to higher algal biomass; but ...
  58. [58]
    Review of the reproductive biology of amphipods and their ...
    Sep 2, 2011 · Spermatophores, sperm transfer, and sperm storage by females. The fertilization process in amphipods in general is poorly understood.
  59. [59]
    [PDF] Energetically costly precopulatory mate guarding in the amphipod ...
    Precopulatory mate guarding (PCMG) is thought to have evolved as a male mating strategy in species in which female receptivity is limited to a short time.Missing: dioecious | Show results with:dioecious
  60. [60]
    Precopulatory Mate Guarding, Mating System and Pairing Parental ...
    Feb 28, 2008 · H. rubra displayed precopulatory mate guarding: males clasped females with their gnathopods during copulation, forming a pair. Males also ...
  61. [61]
    Reproductive strategies of a population of a freshwater amphipod ...
    The aim of this study was to characterize the reproductive strategies of a population of Hyalella bonariensis from southern Brazil.
  62. [62]
    3 Life cycle of amphipod crustaceans. ( A ) Life cycle and direct...
    Adult females carry their embryos in a ventral brood pouch (marsupium; indicated by a square ). The embryonic development takes about 10–12 days at 25 °C (major ...Missing: incubation | Show results with:incubation
  63. [63]
    Hay's Spring Amphipod (Stygobromus hayi) | U.S. Fish & Wildlife ...
    Males and females have been found during collections, but parthenogenesis, a form of asexual reproduction in which embryos develop without fertilization, is ...
  64. [64]
    Reproductive Biology and Egg Production of Hyalella Longistila ...
    Precopulatory mating behavior is the most precise indication that amphipod males are able to reproduce (Borowsky, 1991), while developed ovaries (dark green ...
  65. [65]
    Gammarus tigrinus - Marine Invasions research at SERC
    Males are 8-12 mm in size and females are 6-8 mm. This description is based on Bousfield 1958 and Bousfield 1973. LeCroy (2000) refers to Gulf coast populations ...<|separator|>
  66. [66]
    (PDF) Stages of embryonic development in the amphipod ...
    Aug 9, 2025 · ... Segmentation is apparent. throughout the lateral and dorsal body wall and the. thoracic coxal plates are clearly visible (Fig. 13d). S25 Stage ...
  67. [67]
    [PDF] 503.pdf - Zoological Studies
    In G. aequicauda, as in other amphipods, development is direct, and newly hatched juveniles possess all the structures typical of the adult.
  68. [68]
    Effect of temperature on the duration of egg development, and ...
    The interval between moults is an extension of egg development time, increasing from birth to sexual maturity which is probably reached at instar 6 or 7.
  69. [69]
    Influence of temperature on production of the amphipod Parhyale ...
    Mar 4, 2025 · Artal et al. [27] reported an 89% embryo hatching success rate for P. hawaiensis, a somewhat higher, but comparable rate to that (70–80%) ...
  70. [70]
    stages 1-4 Post-marsupial development of Hyalella pleoacuta ...
    Sep 18, 2009 · In amphipods, fertilization and development of the eggs occur in the marsupium, where the juveniles remain for some time after they hatch ...
  71. [71]
    [PDF] Effect of salinity and temperature on the intra-marsupial ...
    The survival and hatching success are highly dependent on the salinity conditions, while the development time is strongly affected by temperature. High ...
  72. [72]
    The influence of temperature and salinity on the duration of ...
    Results showed that growth was continuous throughout life under all laboratory conditions. Intrinsic growth rates were higher at 20 °C and lower at 10 °C.
  73. [73]
    (PDF) Amphipoda - ResearchGate
    Dec 3, 2014 · G: drawings of post-embryonic stages of hyperiid amphipods referred to as physosoma larvae, lateral views; ( left ) physosoma of Lanceo- lidae; ...
  74. [74]
    Life Cycle and Life History Strategies of Parasitic Crustacea - PMC
    Direct develop larvae (crawl-away larvae)—larval stages that have very low ... Pre-manca—the isopod brood-pouch stage after eyed embryo before manca and is less ...
  75. [75]
    Possible causes of amphi-Atlantic distribution of Orchestia ...
    Jan 9, 2023 · Hypotheses concerning the modern distribution of Orchestia gammarellus (Crustacea, Amphipoda, Talitridae) and its causes in the North Atlantic
  76. [76]
    Could anthropic pressures drive the burrowing behavior of sandy ...
    Aug 8, 2023 · Burying is common among all species inhabiting intertidal beaches, and involves a penetration phase followed by excavation movements, the ...
  77. [77]
    Vertical distribution and daily migrations of hyperiid amphipods in ...
    The vertical distribution and migratory behaviour of hyperiid amphipods were studied in a series of tows carried out during a 48-h sampling period at an ...
  78. [78]
    Drivers of variation in seagrass-associated amphipods across ...
    Amphipods are one of the dominant epifaunal groups in seagrass meadows. However, our understanding of the biogeographical patterns in the distribution of ...
  79. [79]
    Supergiant amphipods from the deep sea - AAAS
    Mar 12, 2012 · Supergiant amphipods are almost 10 times larger than normal ones, found in the deep sea, and are scavengers. The largest recorded was 34 cm ...Missing: benthic epifaunal infaunal
  80. [80]
    Diversity and Zonation of Benthic Amphipod Crustaceans Affected ...
    Amphipod crustacean diversity and zonation are described from the large Mississippi Canyon that extends from the continental shelf across the entire continental ...
  81. [81]
    Themisto amphipods in high-latitude marine pelagic food webs
    Apr 24, 2019 · Themisto are major predators of meso- and macrozooplankton in several major oceanic regions covering shelves to open ocean from the polar regions to the ...Missing: neritic bathyal
  82. [82]
    Salinity modulates thermotolerance, energy metabolism and stress ...
    Nov 17, 2016 · Amphipods from a freshwater population were more sensitive to the thermal challenge, showing higher mortality during acute and gradual ...Missing: gradients bioluminescence
  83. [83]
    Studies on bioluminescent marine amphipods
    May 11, 2009 · The luminescence of a number of amphipods of the genera Scina, Parapronoe, Cyphocaris, Megalanceola, Thoriella and Chevreuxiella/Danaella is ...Missing: abyssal | Show results with:abyssal
  84. [84]
    Global diversity of amphipods (Amphipoda; Crustacea) in freshwater
    There are about 1,870 freshwater amphipod species, with 70% in the Palearctic, 13% Nearctic, 7% Neotropical, 6% Australasian, and 3% Afrotropical regions.Missing: zones | Show results with:zones
  85. [85]
    The giant cryptic amphipod species of the subterranean genus ...
    May 28, 2017 · With more than 350 species described (Horton et al., 2016), the genus Niphargus Schiödte, 1849 is the most species-rich taxon of freshwater ...
  86. [86]
    [PDF] Contribution of the lateral extension of the hyporheic zone in gravel ...
    Aug 2, 2025 · hyporheic invertebrates that inhabit these zones often exhibit specialized adaptations to cope with oxygen stress. 465. These adaptations may ...
  87. [87]
    Invading predatory crustacean Dikerogammarus villosus eliminates ...
    In freshwaters in The Netherlands, we show that the invasive Ponto–Caspian crustacean amphipod Dikerogammarus villosus is rapidly eliminating Gammarus duebeni, ...
  88. [88]
    [PDF] Dikerogammarus villosus ERSS - U.S. Fish and Wildlife Service
    The Killer Shrimp (Dikerogammarus villosus) is a small freshwater and brackish amphipod native to the Ponto-Caspian basin of Europe. The species can tolerate a ...
  89. [89]
    (PDF) Biology of Terrestrial Amphipods - ResearchGate
    Aug 6, 2025 · The floor of mesic forests in the Southern Hemisphere and the tropics is the major habitat type in which landhoppers (Talitridae) are found.
  90. [90]
    Terrestrial Environments | Evolution and Biogeography: Volume 8
    Among all these crustacean groups, only the family Talitridae among the Amphipoda and the suborder Oniscidea among the Isopoda show full adaptations to live on ...Missing: jumping | Show results with:jumping
  91. [91]
    Dietary analysis of the marine Amphipoda (Crustacea
    Amphipods inhabit a variety of marine environments and, in consequence, they show a high diversity of feeding habits; they can feed on debris, detritus, ...
  92. [92]
    Feeding habits of amphipods (Crustacea: Malacostraca) from ...
    The present work is the first comprehensive study on the feeding habits of amphipod species inhabiting soft bottom communities.
  93. [93]
    Extreme food webs: Foraging strategies and diets of scavenging ...
    Jan 17, 2007 · The extreme foraging flexibility of scavenging amphipods may ultimately contribute to their success in severely food- limited hadal ecosystems. ...
  94. [94]
    Comparative functional morphology of feeding in three species of ...
    It was found that Caprella penantis Leach fed primarily by filter-feeding and scraping; Paracaprella tenuis Mayer relied on filter-feeding, scraping, scavenging ...
  95. [95]
    (PDF) Feeding habits of caprellids (Crustacea: Amphipoda) from the ...
    Aug 5, 2025 · The present study reveals that caprellids from the Pacific coast of Mexico are able to feed on hydroids but not on gorgonians.
  96. [96]
    DNA metabarcoding reveals a diverse, omnivorous diet of Arctic ...
    Feb 13, 2024 · In this study, we aimed to verify the role of jellyfish and other metazoans as food sources in the Arctic polar night food web, when pelagic resources are ...
  97. [97]
    Artificial light at night alters foraging behavior of freshwater ...
    Oct 3, 2024 · Artificial light at night alters foraging behavior of freshwater amphipods depending on the light spectrum and the presence of predation cues.Missing: swarming symbiotic
  98. [98]
    (PDF) Trophic diversity in amphipods within a temperate eelgrass ...
    Aug 10, 2025 · Stable isotope data confirmed that epiphytes, detritus, and mesozooplankton fragments were major food sources for amphipods in the eelgrass bed.
  99. [99]
    Lake trout–induced spatial variation in the benthic invertebrates of ...
    Lake trout may alter the biomass and body size of amphipods because these fish also feed on invertebrates. Amphipods comprised 25% of the diet of juvenile ...
  100. [100]
    (PDF) Feeding ecology and predation impact of the recently ...
    Stomach content analyses showed that these amphipods consumed chiefly copepods, in particular, the copepodite stages CIV and CV of Calanus finmarchicus.
  101. [101]
    The role of hyperiid parasites as a trophic link between jellyfish and ...
    May 19, 2025 · In turn, hyperiid amphipods parasitising jellyfish can occur in such great numbers that they represent an important trophic link to diverse ...Missing: hooks | Show results with:hooks<|separator|>
  102. [102]
    High parasite diversity in the amphipod Gammarus lacustris in a ...
    Oct 5, 2020 · Amphipods are often key species in aquatic food webs due to their functional roles in the ecosystem and as intermediate hosts for trophically ...
  103. [103]
    Gammaridae - an overview | ScienceDirect Topics
    Gammaridae is a family of amphipods, with diverse Gammarus species, that are important in freshwater ecosystems and sensitive to chemical stressors.
  104. [104]
    Potential impact of invasive amphipods on leaf litter recycling in ...
    Aug 5, 2025 · Furthermore, amphipods are frequently used as bioindicators of pollution and other environmental stressors in a variety of aquatic ecosystems.
  105. [105]
    Gammarus Spp. In Aquatic Ecotoxicology and Water Quality ...
    This review summarizes the vast number of studies conducted with Gammarus spp. for evaluating aquatic ecotoxicology endpoints.Missing: bioindicator | Show results with:bioindicator
  106. [106]
    Studies of Commensal Leucothoid Amphipods: Two New Sponge ...
    Aug 7, 2025 · These amphipods feed by filtering particles from the water passing through the central cavity of the sponge, trapping them on the numerous setae ...
  107. [107]
    Unveiling the complexity and ecological function of aquatic ...
    Other mutualistic interactions can be observed between mesograzers, such as amphipods, and macroalgae. Amphipods facilitate macroalgae by feeding on smaller ...
  108. [108]
    Echinogammarus ischnus - Marine Invasions research at SERC
    It has a short, triangular rostrum. The eyes are kidney-shaped and twice as long as wide. Antennae 1 and 2 are roughly equal in length. Antenna 1 has a ...
  109. [109]
    [PDF] 2020 UPDATE TO “AN IMPACT ASSESSMENT OF GREAT LAKES ...
    Dec 21, 2021 · We estimate 17% (previously 14 to 16%) of the nonindigenous species found in the Great Lakes have moderate to high socioeconomic impact. Table ...
  110. [110]
    First Mesozoic amphipod crustacean from the Lower Cretaceous of ...
    The first Mesozoic amphipod, Gammaroidorum vonki, found in the Early Cretaceous Wealden of the Weald, is morphologically somewhat similar to a diminutive ...
  111. [111]
    [PDF] Further records of Amphipoda from Baltic Eocene amber with first ...
    Feb 20, 2014 · In the present paper we provide a description of two new amphipod species of the genera Palaeogammarus and Synurella found in Baltic Eocene ...
  112. [112]
    A new terrestrial amphipod from tertiary amber deposits of Chiapas ...
    Jan 10, 2009 · This new taxon is the first known fossil member of the amphipod family Talitridae and superfamily Talitroidea. It appears intermediate between ...
  113. [113]
    A new fossil talitrid amphipod from the lower early Miocene Chiapas ...
    A review of the amphipod fossil record is given which demonstrates that the amphipods first appeared as fossils in the Eocene.
  114. [114]
    Evolutionary morphology of the circulatory system in Peracarida ...
    Mar 5, 2010 · Over the years various phylogenetic studies have supported a sister-group relationship between Amphipoda and Isopoda (e.g. Schram, 1986 ...<|control11|><|separator|>
  115. [115]
    SPECIAL SECTION ON THE BIOLOGY OF PERACARIDS
    Jun 3, 2016 · Peracarida is generally divided into 12 accepted or- ders: Amphipoda Latreille, 1816 (around 10,000 described species), Bochusacea Gutu and ...
  116. [116]
    Molecular phylogenetic evidence for the reorganization of the ...
    The hyperiid amphipods are classically defined as an exclusively pelagic group broadly inhabiting oceanic midwater environments and often having close ...
  117. [117]
    The late blooming amphipods: global change promoted post-Jurassic ecological radiation despite Palaeozoic origin
    **Summary of Phylogenetic Relationships and Divergence Times in Amphipoda:**
  118. [118]
    Brooding and embryonic development in the crustacean ...
    Known embryological development for P. formica includes three post gastrulation stages. In the current study, brooding and embryological development in this ...
  119. [119]
    [PDF] AN ALTERNATIVE PHYLOGENY OF PERACARID CRUSTACEANS ...
    Two lineages are immediately apparent: 1, a heterogeneous line con- taining the Amphipoda, Mysidacea, and Thermosbaenacea; and 2, a homogeneous "mancoid" line ...
  120. [120]
    Peracarid monophyly and interordinal phylogeny inferred from ...
    Apr 1, 2005 · Peracarids are a large group of malacostracan crustaceans whose systematics and phylogeny are uncertain. The present phylogenetic study of ...<|control11|><|separator|>
  121. [121]
    Major Revisions in Pancrustacean Phylogeny and Evidence of ...
    Aug 8, 2023 · Our MCMCtree divergence time estimates retrieved deep splits of arthropods and the three main pancrustacean clades (Oligostraca, Allotriocarida, ...
  122. [122]
    Use of Gammarus pulex (L.) in safety evaluation tests
    A simple and effective method for the laboratory culture of Gammarus pulex is described. Over 70% of juveniles survived, reaching sexual maturity within 130 ...
  123. [123]
    (PDF) Gammarus fossarum as European amphipod species for ...
    Oct 20, 2025 · The protocol assesses toxicity of persistent chemicals by measuring the effect of spiked sediment on adults' reproduction. The test guideline ...
  124. [124]
    Marine amphipods (Parhyale hawaiensis) as an alternative feed for ...
    Oct 19, 2021 · The present study assessed the nutritional value of an interesting marine amphipod (Parhyale hawaiensis), and evaluates through a feeding trial ...
  125. [125]
    Nutritional value of the amphipod Bemlos quadrimanus sp. grown in ...
    Feb 25, 2021 · This study demonstrates that amphipod B. quadrimanus provide an important source of protein that shows potential to replace fishmeal in shrimp feed.Missing: Hyperia | Show results with:Hyperia
  126. [126]
    Effects of culture system and diet on amphipod (Parhyale ... - Nature
    Aug 4, 2025 · Results of this study suggest that utilization of a biofloc culture system may both improve water quality and increase P. hawaiensis production.
  127. [127]
    A psychrophilic caseinolytic aspartic protease from the freshwater ...
    In the present study, an aspartic protease is purified from the freshwater crustacean amphipod Gammarus bakhteyaricus (Khalaji-Pirbalouty & Sari, 2004). This ...
  128. [128]
    Polysaccharide hydrolase of the hadal zone amphipods Hirondellea ...
    We found that Hirondellea gigas thrived in the Challenger Deep possessed polysaccharide hydrolases as digestive enzymes. To obtain various enzymes of other H.
  129. [129]
    Report Nanostructures and Monolayers of Spheres Reduce Surface ...
    Nov 21, 2016 · We investigated the cuticle surfaces of seven species of hyperiids (Crustacea; Amphipoda) ... anti-fouling or prey capture. We also acknowledge ...
  130. [130]
    Chitosan from Marine Amphipods Inhibits the Wilt Banana Pathogen ...
    Nov 17, 2023 · In this work, we extracted chitosan from marine amphipods associated with aquaculture facilities and tested its use in crop protection.
  131. [131]
    First evidence of microplastics ingestion in benthic amphipods from ...
    Aug 7, 2025 · The present paper provides the first record of ingestion of microplastics in natural context by Gammarus setosus from Svalbard Archipelago.
  132. [132]
    Microplastics and synthetic particles ingested by deep-sea ...
    Here, we detect the presence of ingested microplastics in the hindguts of Lysianassoidea amphipod populations, in six deep ocean trenches from around the ...Missing: sentinels | Show results with:sentinels
  133. [133]
    Differential effects of microplastic exposure on leaf shredding rates ...
    Oct 13, 2023 · This study investigates the effect of microplastic exposure on leaf consumption by two Gammarus species in Ireland, the native Gammarus duebeni celticus, and ...
  134. [134]
    Microplastic pollution in the food web: observation of ingestion by ...
    Jan 9, 2023 · Several studies have been carried out on the ingestion of microplastics by Amphipods, both in laboratory conditions (Blarer & Burkhardt-Holm ...
  135. [135]
    [PDF] Gammarus spp. in Aquatic Ecotoxicology and Water Quality ...
    OECD-proposed test species, because Gammarus spp. naturally occur in streams of the Northern Hemisphere. The breadth of ecotoxicological studies published on.
  136. [136]
    Skeleton Shrimp (Caprella spp.) | Live Maine Amphipods for Study
    In stockUses: Excellent for aquariums, live plankton demonstrations, marine invertebrate study, and behavioral observation. Note: This is a live marine specimen.Missing: commercial ornamental
  137. [137]
    The Invasive Amphipod Dikerogammarus villosus Impacts Detritus ...
    Jun 7, 2023 · This study investigated how these stressors interact to impact the vital ecosystem process of shredding by keystone species in temperate freshwater ecosystems.
  138. [138]
    Seasonal changes in biodiversity of native and non-native ...
    Jul 4, 2024 · In this study, we conducted a comparative assessment of seasonal variations in amphipod communities at three southeastern Baltic Sea locations.
  139. [139]
    Amphipods are Good Bioindicators of the Impact of Oil Spills on Soft ...
    It is suggested to focus monitoring after a spill only on a single amphipod group proposed as a bioindicator for detecting the impact of pollution.Missing: pollution | Show results with:pollution
  140. [140]
    [PDF] Using amphipods as bioindicators of metal pollution in the marine ...
    Amphipods are internationally used as a biomonitor and bioindicator species as a means of assessing the natural environment and changes in response to pollution ...
  141. [141]
    Freshwater amphipods (Gammarus pulex/fossarum) and brown trout ...
    Aug 24, 2020 · Freshwater amphipods would appear to potentially serve as bioindicators for PFC contamination in small streams. These organisms are ubiquitous ...
  142. [142]
  143. [143]
    [PDF] Options for the control of Dikerogammarus villosus (killer shrimp ...
    Control options include technological methods (pheromone, RNAi, biotechnologies) and management strategies like eradication, reproductive removal, containment, ...
  144. [144]
    Management Strategies: Killer Shrimp, Dikerogammarus villosus ...
    Apr 13, 2018 · So, most management strategies for invasive amphipods are not designed to eradicate, but to control. However, in the USA and Canada, a law ...
  145. [145]
    Amphipod crustaceans and environmental disturbance: a review
    This review of the literature shows that amphipod crustaceans create disturbance through predation on commercial harvests, attacks on macroalgae, introductions ...