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Diel vertical migration

Diel vertical migration (DVM), also known as diurnal vertical migration, is a synchronized daily movement pattern observed in aquatic environments, where planktonic and nektonic organisms—such as zooplankton, copepods, euphausiids, fish, and cephalopods—ascend to surface waters at dusk to feed and descend to deeper layers before dawn, often covering distances from hundreds of meters to over 800 meters in the water column.^[1] This behavior is ubiquitous across marine and freshwater systems, from epipelagic to bathypelagic zones, and represents the largest biomass migration on Earth, involving billions of individuals globally.^[2] The migration is primarily driven by a balance between foraging opportunities and predation risk, with organisms exploiting nutrient-rich surface layers under the cover of darkness while retreating to deeper, darker depths during daylight to evade visually hunting predators.^[3] Light intensity serves as the key environmental cue, often aligning with specific isolumes (e.g., around 10⁻³ W m⁻²), though internal circadian rhythms, temperature gradients, prey availability, and oxygen levels can modulate the timing and amplitude of movements.^[3] In some cases, such as with certain dinoflagellates or deep-sea plankton, migrations may also optimize reproduction or reduce exposure to harmful ultraviolet radiation.^[1]^[2] Ecologically, DVM profoundly influences aquatic food webs and biogeochemical cycles, comprising 40–60% of mesozooplankton biomass in many ocean regions and forming detectable deep-scattering layers via sonar.^[3] By transporting organic carbon from surface productivity to the mesopelagic zone through active metabolism and fecal pellets, it enhances the biological pump, contributing 15–40% to particle export and up to 50% of respiration in migration layers, thereby playing a critical role in global carbon sequestration and nutrient recycling.^[3] This process also generates turbulence, affects primary production, and supports higher trophic levels, underscoring DVM's integral role in maintaining ocean ecosystem dynamics and influencing climate regulation.^[2]

Introduction

Definition and scope

Diel vertical migration (DVM) refers to the synchronized daily vertical displacement of aquatic organisms, in which they typically ascend from deeper waters to surface layers at dusk and descend back to depth at dawn, following a 24-hour cycle tightly linked to the light-dark transition.^[3] This behavior represents a fundamental adaptive strategy in aquatic ecosystems, enabling organisms to optimize foraging and survival amid varying environmental conditions.^[4] The scope of DVM encompasses primarily planktonic and nektonic organisms, including zooplankton such as copepods and krill, micronekton like small fish and mysids, and various larval stages.^[3] It is distinct from random or irregular vertical movements, as DVM exhibits precise, rhythmic patterns driven by predictable environmental cycles rather than stochastic factors.^[4] This phenomenon is observed across marine and freshwater systems, though it is most pronounced in oceanic environments where light penetration structures the water column.^[3] Basic patterns of DVM include migration amplitudes ranging from 100 to 800 meters in oceanic settings, with organisms often phase-locked to the solar cycle—ascending shortly before sunset and descending before sunrise to align with optimal light conditions.^[3] The behavior is highly ubiquitous, affecting the vast majority of zooplankton species and, in some systems, a substantial portion of the zooplankton biomass participates in these migrations.^[4] In terms of ecological scale, DVM constitutes one of the largest biomass migrations on Earth, involving an estimated ~5 billion tons of organisms and dwarfing all terrestrial animal migrations combined.^[5]

Global occurrence and key examples

Diel vertical migration (DVM) is a ubiquitous phenomenon observed across diverse aquatic ecosystems worldwide, encompassing marine, freshwater, and estuarine environments, and involving a broad array of organisms from zooplankton to fish. In marine settings, it is particularly prominent in the open ocean, where vast populations of zooplankton and micronekton undertake synchronized daily movements spanning hundreds of meters. For instance, Antarctic krill (Euphausia superba) in the Southern Ocean perform extensive DVM, descending to depths of 200–400 meters during the day and ascending to surface waters at night, facilitating nutrient transport and predator avoidance in these polar regions.^[6] Similarly, mesopelagic fishes such as lanternfish (family Myctophidae) dominate the deep scattering layers of the world's oceans, migrating from depths of 500–750 meters by day to the upper 200 meters at night, contributing significantly to the global biomass flux in the mesopelagic zone.^[7] In freshwater systems, DVM is equally widespread, especially among cladocerans in temperate lakes, where it helps regulate predator-prey dynamics in stratified water columns. A key example is the water flea Daphnia species, which commonly migrate vertically by 5–20 meters daily, residing in deeper, cooler hypolimnetic layers during daylight to evade visual predators like fish and rising to the epilimnion at night for foraging on phytoplankton.^[8] This behavior is documented in numerous lakes across North America and Europe, underscoring its role in maintaining ecosystem balance in these enclosed basins and driving vertical transport of organic matter at ecosystem scales.^[9] Estuarine and coastal habitats exhibit more variable DVM patterns, often modified by tidal influences and salinity gradients in shallow bays. Mysids (Crustacea: Mysidae), such as Mysis diluviana, demonstrate partial DVM in these dynamic environments, with only a fraction of the population migrating from benthic substrates to the water column at night, while others remain in place, allowing niche partitioning and adaptation to fluctuating conditions.^[10] This partial migration is observed in coastal systems like those around the Great Lakes and North American estuaries, where it supports energy transfer between seafloor and surface layers.^[11] The scale of DVM is immense, representing one of the planet's largest synchronized animal movements by biomass, with global oceanic fluxes estimated to rival terrestrial migrations in magnitude and involving trillions of individuals nightly. Recent studies up to 2025 highlight ongoing observations of DVM in changing environments; for example, in subarctic epipelagic waters off Greenland, capelin (Mallotus villosus) and cod (Gadus morhua) exhibit pronounced diel movements that structure feeding interactions within the water column.^[12] Likewise, larval and juvenile small yellow croaker (Larimichthys polyactis) in coastal waters of the Yangtze River Estuary, China, perform DVM to optimize transport and survival, ascending at night to exploit faster currents for offshore dispersal.^[13] These examples illustrate the adaptability and ecological significance of DVM across global aquatic realms.

Historical discovery

Early observations

The earliest documented observations of diel vertical migration emerged in the 19th century, primarily through the work of naturalists employing rudimentary sampling techniques in both freshwater and marine environments. In 1817, French naturalist Georges Cuvier provided the first description of the phenomenon in the freshwater cladoceran Daphnia, noting its retreat to deeper waters at midday to avoid intense light and its ascent toward the surface in the evening as light diminished.^[14] This observation highlighted a rhythmic, light-driven behavior, though it was initially confined to shallow lakes and lacked broader ecological context. Subsequent marine explorations built on such anecdotal records, shifting focus to oceanic plankton. Marine records began with depth-stratified net sampling during mid-century expeditions. In the 1850s, British naturalist Edward Forbes, during his expeditions in the Mediterranean Sea aboard vessels like HMS Beacon, used dredges and tow nets to collect plankton and benthic organisms, observing progressive changes in species abundance and composition with depth, emphasizing zonation patterns where surface-dwelling plankton appeared scarcer during daylight hours compared to nighttime hauls, though he attributed variations partly to tidal influences rather than strictly diel cycles.^[14] These qualitative insights marked an initial recognition of vertical structuring in Mediterranean plankton communities, influencing later oceanographic surveys. Systematic evidence from global voyages further illuminated the phenomenon. The HMS Challenger expedition (1872–1876), a landmark British scientific circumnavigation, conducted repeated plankton tows at varying depths and times across the Atlantic, Pacific, and Indian Oceans, documenting diurnal variations in catches from deep-sea hauls—such as increased surface abundance of crustaceans and radiolarians at night.^[14] Reports by expedition naturalists, including John Murray, noted these patterns in open-ocean settings, linking them to light availability and providing the first global-scale hints of widespread diel migration among pelagic organisms.^[15] Concurrently, late-19th-century fisheries observations offered practical corroboration; for instance, 1880s observations from Scottish herring fisheries, such as those by Brook (1886) in Loch Fyne, noted daytime near-surface aggregations of copepods like Calanus finmarchicus that decreased with herring presence, suggesting behavioral responses to predation exploited by fishermen but initially unexplained beyond seasonal cues.^[15] Despite these pioneering efforts, early methods imposed significant limitations on understanding. Reliance on manual net tows—often coarse-meshed and deployed sporadically—yielded inconsistent data, capturing only discrete depth intervals and missing the continuous, real-time dynamics of migration.^[14] Such techniques, as used by Forbes and Challenger scientists, frequently confounded diel patterns with tidal or weather effects, resulting in incomplete profiles until the advent of more precise, continuous sampling in the 20th century. These foundational observations nonetheless established diel vertical migration as a recurrent ecological feature, later validated through acoustic and optical technologies.^[14]

Scientific advancements and key researchers

In the early 20th century, significant methodological progress in studying diel vertical migration (DVM) came from the development of quantitative sampling tools, notably the Clarke-Bumpus plankton sampler introduced in the 1930s by George L. Clarke. This device allowed for precise, depth-specific collections of plankton, enabling researchers to document vertical distributions and migration patterns with greater accuracy than previous net hauls, thus providing foundational quantitative data on DVM amplitudes in oceanic waters.^[16]^[17] Clarke, a pioneering oceanographer, advanced quantitative sampling and hypotheses on light-driven migrations during the 1930s, such as the rate-of-change hypothesis linking DVM to irradiance variations, marking a shift from descriptive to mechanistic observations of subsurface behaviors.^[17] A major breakthrough occurred during World War II when sonar operators detected moving 'false bottoms' in the ocean, later identified as deep scattering layers (DSLs) formed by migrating zooplankton and fish, providing the first acoustic evidence of widespread DVM.^[18] Building on this, the mid-20th century saw John E. Harris's influential work in the 1950s and 1960s, where he demonstrated the role of endogenous rhythms in driving DVM through laboratory experiments on copepods, isolating circadian clocks from external cues like light.^[19]^[14] Harris's 1963 paper established that internal biological timers persist under constant conditions, solidifying the endogenous component of migration control.^[19] By the 1960s, electronic depth sounders enhanced resolution of migration dynamics, revealing daily vertical excursions of up to several hundred meters in zooplankton layers, which previous sampling had underestimated.^[17] The advent of acoustic Doppler current profilers (ADCPs) in the 2000s revolutionized real-time tracking, allowing non-invasive monitoring of DVM in situ across large water columns by detecting acoustic backscatter from migrating organisms.^[20]^[21] Milestone expeditions like the Tara Oceans voyage (2009–2013) integrated multidisciplinary sampling to map global DVM patterns, combining net tows, acoustics, and imaging to quantify vertical fluxes and community structures in the upper ocean.^[22] Recent modeling efforts in the 2020s, such as the 2025 study by Vilain et al., which used Continuous Plankton Recorder data and Map Equation clustering to show how DVM influences copepod bioregions in the North Atlantic, linking behavioral patterns to large-scale biogeographic boundaries.^[23] Technological evolution has progressed from discrete net-based sampling to advanced satellite telemetry, which tracks individual migrators' depths via pop-up tags, and genomic sequencing, revealing genes associated with light sensitivity and circadian regulation in DVM performers.^[24]^[25] These innovations, exemplified by high-throughput eDNA analysis, now enable identification of migration-related genetic adaptations across taxa.^[25]

Types of vertical migration

Diel patterns

Diel vertical migration primarily manifests in three distinct patterns among aquatic organisms, particularly zooplankton: nocturnal, reverse, and twilight migrations. Nocturnal migration represents the standard and most prevalent subtype, in which organisms ascend from deeper waters to the surface layers around dusk, remain in the upper water column throughout the night, and descend to deeper depths by dawn. This behavior is exhibited by the majority of zooplankton species in marine environments, including many copepods and euphausiids.^[3]^[26] For instance, nearly all mesopelagic zooplankton engage in this pattern, contributing to massive daily biomass displacements across ocean basins.^[26] Reverse migration, a less common variant, involves organisms rising to shallower depths during the day and sinking to deeper layers at night. This subtype is observed in specific taxa, such as juveniles or males in certain copepod species, and in some freshwater cladocerans.^[14]^[27] Examples include smaller individuals of Pseudocalanus spp. in coastal bays, which follow this inverted cycle relative to the dominant nocturnal pattern.^[14] Twilight migration features limited vertical excursions synchronized with dawn and dusk transitions, typically involving partial ascents and descents confined to shallow depths with amplitudes under 50 m. This pattern predominates in tidally influenced coastal regions, such as among mysid crustaceans like Mysis spp., where movements remain relatively shallow and do not span the full water column.^[1]^[28] Overall, nocturnal migration dominates oceanic systems, while reverse and twilight forms are more regionally specific, with the latter often tied to dynamic nearshore habitats.^[29] These diel patterns may exhibit seasonal variations in amplitude or timing.^[14]

Non-diel patterns

Non-diel vertical migrations encompass patterns that operate on timescales longer than the daily cycle, including seasonal and ontogenetic shifts in depth distribution among zooplankton. Seasonal vertical migration (SVM) involves gradual depth changes over months, often driven by annual environmental variations, where organisms descend to deeper waters during unfavorable periods such as winter for overwintering. For instance, large polar copepods like Calanus finmarchicus and C. glacialis exhibit SVM with amplitudes reaching over 1000 m in polar seas, relocating from surface layers in summer to depths exceeding 500 m in winter to conserve energy in dormancy.^[30]^[31] Ontogenetic vertical migration (OVM) refers to progressive depth shifts across an individual's lifespan, correlating with developmental stages and body size increases. In calanoid copepods such as Eucalanus inermis, naupliar stages remain near the surface to access phytoplankton, while copepodite and adult stages descend deeper, often into oxygen minimum zones below 200 m, with deepening continuing through maturation.^[32]^[33] This pattern is prevalent in many long-lived zooplankton species, including approximately half of calanoid copepod taxa, where OVM facilitates stage-specific resource utilization and predator avoidance.^[34] These non-diel patterns interact with diel vertical migration by establishing baseline depths that modulate daily amplitudes; for example, seasonal overwintering descent in polar copepods reduces the scope for diel excursions during winter, while ontogenetic deepening in juveniles limits their participation in full adult diel cycles. Body size effects, linked to ontogeny, further influence these shifts by affecting metabolic demands and swimming capacity.^[30]^[31] Such interactions highlight how extended migrations provide a framework for daily behaviors in diverse aquatic ecosystems.

Controls on diel migration

Endogenous mechanisms

Endogenous mechanisms underlying diel vertical migration (DVM) primarily involve internal biological clocks that drive and synchronize migratory behaviors, independent of immediate environmental fluctuations. These circadian rhythms, with free-running periods typically ranging from 23 to 25 hours, enable organisms to anticipate daily cycles and initiate vertical movements even in constant conditions. In the freshwater cladoceran Daphnia species, for instance, these rhythms persist under constant darkness, maintaining synchronized ascent and descent patterns that mirror natural DVM. Similarly, in marine copepods like Calanus finmarchicus, laboratory experiments demonstrate endogenous oscillations in swimming activity and respiration that continue for several days in the absence of light-dark cues, confirming the role of an internal timing mechanism.^[35] At the molecular level, these rhythms are governed by circadian clock genes, such as period (per) and cryptochrome (cry), which form transcriptional-translational feedback loops in crustaceans. These loops oscillate gene expression to regulate physiological processes, including buoyancy adjustments and swimming behaviors critical for DVM. In Antarctic krill (Euphausia superba), daily peaks in per and cry expression align with migratory phases, driving metabolic and locomotor changes that facilitate vertical positioning.^[36] Orthologs of these genes, conserved across arthropods, have been identified in the genome of Daphnia pulex, underscoring their ancient role in synchronizing crustacean behaviors.^[37] Body size influences the expression of these endogenous mechanisms through metabolic scaling, where larger individuals often exhibit greater migration amplitudes or deeper depths due to allometric relationships. For example, larger copepods descend to greater depths during daylight, as observed in field studies.^[38] This arises from endogenous programming that adjusts vertical excursions to optimize individual fitness.^[39] Supporting evidence for these internal drivers comes from controlled laboratory studies and genetic manipulations. Experiments in perpetual darkness reveal rhythm persistence, as seen in ostracods (Asterope and Philomedes) where circadian swimming patterns endure without external zeitgebers.^[40] Furthermore, genetic knockouts of clock genes disrupt DVM timing; a 2024 study in Daphnia showed that per gene inactivation abolishes sustained vertical migration under constant conditions, highlighting the clock's essential role in rhythm generation.^[41] These findings collectively affirm that endogenous clocks provide a robust, heritable framework for DVM across zooplankton taxa.

Exogenous cues

Light serves as the primary exogenous cue, or zeitgeber, regulating the timing of diel vertical migration in most aquatic organisms, with changes in intensity and spectral quality triggering ascent and descent phases. At dawn, increasing blue light wavelengths (approximately 450-500 nm) prompt descent to deeper waters, enabling organisms to avoid visually hunting predators during daylight hours, while at dusk, red-shifted light facilitates ascent to surface layers for feeding.^[42] This response is highly sensitive, with migration often initiating at low light intensities around 10^{-7} μmol photons m^{-2} s^{-1} for species such as Calanus and krill, with thresholds varying from 10^{-8} to 10^{-6} μmol photons m^{-2} s^{-1} across taxa and habitats.^[43] Temperature gradients, particularly thermal stratification in the water column, modulate the speed and extent of migration, often accelerating movements in response to warmer surface layers. In stratified environments, organisms like copepods exhibit faster ascent rates in epilimnetic waters (typically 5-15°C warmer than deeper layers), which can enhance overall migration efficiency by reducing exposure time in vulnerable zones.^[44] For instance, in species such as Calanus finmarchicus, warmer temperatures increase swimming velocities, allowing quicker transitions between depths.^[44] Additional exogenous factors include salinity gradients in estuarine systems, which guide orientation and vertical positioning during migration. In coastal zones with sharp haloclines, lower salinity in surface waters can cue upward movements, helping larvae and juveniles maintain position against tidal flows.^[45] Some crustaceans sense hydrostatic pressure changes, providing depth feedback that fine-tunes descent depths and prevents overshooting. Chemical cues from predators, known as kairomones, further enhance descent behaviors, particularly in freshwater and coastal zooplankton like Daphnia, where these infochemicals induce deeper positioning even under low light conditions to evade detection.^[46] In coastal areas, tidal cycles synchronize migrations, with flood tides often aligning with ascent to exploit nutrient-rich inflows.^[47] Interactions among these cues emphasize light's dominance, yet temperature can amplify its effects; for example, thermal gradients of 5-10°C across the water column have been observed to roughly double migration rates in dinoflagellates and copepods by boosting metabolic and swimming responses to light signals.^[48] Such synergies ensure adaptive precision in dynamic aquatic environments.

Adaptive functions

Diel vertical migration (DVM) primarily functions to minimize encounters with visually oriented predators, such as planktivorous fish, by exploiting variations in light availability across depths. During daylight hours, migrating zooplankton and nekton descend to depths exceeding 200 meters, where light penetration is limited and visual foraging by predators is severely impaired, thereby reducing predation risk in the well-lit surface layers. At night, ascent to shallower waters occurs under conditions of low ambient light, further limiting the effectiveness of sight-based predation. This temporal and spatial separation is a key adaptive strategy observed across diverse aquatic systems, from freshwater lakes to oceanic environments.^[3]^[49] In addition to light-mediated behaviors, chemical signals known as kairomones—released by predators like fish—play a crucial role in enhancing DVM as a predator avoidance mechanism, particularly in species such as Daphnia. These infochemicals induce rapid and intensified migratory responses, prompting individuals to increase the depth or amplitude of their descent during the day, which strengthens separation from predators. For instance, the bile salt 5α-cyprinol sulfate, a specific kairomone from fish, elicits significant daytime downward shifts in Daphnia at concentrations as low as 100 pM, amplifying the behavioral response beyond light cues alone. This chemical induction allows prey to fine-tune their migration based on perceived predation threat, often resulting in more pronounced avoidance patterns.^[50]^[51] While DVM provides clear anti-predator benefits, it incurs metabolic costs associated with active swimming, which can elevate respiration rates and consume a substantial portion of the daily energy budget in some zooplankton. These costs are offset by substantial survival advantages; modeling studies indicate that DVM can substantially reduce predatory mortality when visual predators dominate, far outweighing the energetic expenditure. Field and enclosure experiments further demonstrate that non-migrating individuals experience markedly higher predation rates than migrants, due to prolonged exposure in vulnerable surface waters. Reverse DVM patterns, where prey ascend during the day and descend at night, also emerge in systems dominated by non-visual invertebrate predators, such as chaetognaths, to minimize overlap with these nocturnal hunters and protect vulnerable life stages.^[3]^[52]

Physiological and foraging advantages

Diel vertical migration provides significant physiological benefits to migrating organisms, particularly through metabolic optimization enabled by thermal stratification in aquatic environments. By ascending to warmer surface waters at night for feeding and descending to cooler deeper layers during the day, zooplankton such as copepods and daphnids experience reduced respiration rates in the colder depths, leveraging the Q₁₀ effect where metabolic rates typically decrease by a factor of 2–3 for every 10°C drop in temperature. For instance, at 10°C compared to 20°C, metabolic rates can be 2–3 times lower, allowing energy that would otherwise be expended on maintenance to be allocated toward growth and reproduction.^[53] This strategy can yield significant daily energy savings relative to constant surface residency, assuming equal time spent in warm and cold layers and a Q₁₀ of 2, as modeled in early bioenergetic analyses. Foraging advantages further enhance the adaptive value of diel migration, as nighttime surface excursions grant access to concentrated phytoplankton and microzooplankton resources that are scarce in deeper waters. In many systems, primary production is confined to the euphotic zone, enabling migrators like calanoid copepods to ingest substantially more carbon during brief surface sojourns compared to what would be available at depth. This pulsed feeding optimizes energy intake while minimizing time in risky surface conditions, supporting higher growth and reproductive output over non-migratory conspecifics.^[54] Migration also mitigates physiological stress from ultraviolet (UV) radiation, a potent DNA-damaging agent in sunlit surface layers. Daytime descent to depths below 10 m in clear waters reduces UVB exposure by approximately 99%, as attenuation in even oligotrophic conditions limits penetrable UV to shallow euphotic zones. Some species, including daphnids, further adapt through inducible pigmentation changes, such as increased melanin production, to shield against residual UV during ascent or in shallower migrations.^[54] In clear oligotrophic waters, where visual foraging challenges arise due to high transparency, diel migration enhances feeding efficiency by timing surface access to nocturnal low-light periods, when reduced visibility aids prey capture without compromising the benefits of deeper daytime refuge. This transparency-driven pattern allows migrators to exploit sparse surface resources more effectively than in turbid systems, where such vertical shifts are less pronounced.

Ecological roles

Carbon cycling and biological pump

Diel vertical migration (DVM) contributes to the ocean's biological pump through active transport of carbon, where migrating organisms, primarily zooplankton and micronekton, feed on phytoplankton and particulate organic carbon in surface waters at night before descending to deeper layers during the day, where they respire carbon dioxide, excrete dissolved organic carbon, and produce fast-sinking fecal pellets. This process decouples grazing from remineralization, effectively exporting carbon from the euphotic zone to the mesopelagic, with active flux via DVM estimated to account for 15–20% of passive particulate organic carbon export in many regions.^[55] Globally, DVM-mediated carbon export is quantified at approximately 0.5–1.3 Gt C year⁻¹, primarily through respiration, fecal pellet production, and excretion, representing about 10% of total oceanic carbon export. Recent 2025 modeling of micronekton DVM, using trait-based approaches that account for size-dependent production rates, indicates these organisms contribute 10–18% to total active flux relative to passive sinking, with fishes showing the highest efficiency due to deeper migrations and rapid gut processing, particularly in temperate regions during summer.^[56]^[57] DVM amplifies the biological pump by enhancing overall carbon sequestration; one-dimensional models demonstrate that migration-induced decoupling of feeding and remineralization increases deep export by approximately 15% compared to non-migratory scenarios, with twilight zones (100–200 m depths) acting as hotspots where up to 50% of total respiration occurs due to concentrated migrant biomass.^[58] This active component supplements passive particle sinking, elevating net flux to the deep ocean and supporting long-term carbon storage. Empirical evidence from sediment trap deployments correlates DVM amplitude with elevated carbon export, as traps capture increased fecal pellet fluxes during periods of high migration intensity, often exceeding passive sinking rates. In the North Atlantic, studies on copepod DVM, particularly of Calanus finmarchicus, reveal seasonal lipid storage and diapause migrations export 2–6 g C m⁻² year⁻¹ via active transport, comparable to sinking fluxes at 600–1,400 m depths and verified against trap data showing repackaged primary production in pellets.

Food web dynamics and biogeography

Diel vertical migration (DVM) plays a pivotal role in structuring aquatic food webs by facilitating the transfer of energy and nutrients between surface and deeper layers, effectively linking epipelagic primary production to mesopelagic consumers.^[59] Migrating zooplankton and micronekton, such as euphausiids, ascend to the surface at night to feed on phytoplankton and then descend during the day, exporting organic matter downward through respiration, excretion, and predation mortality.^[59] This process supports higher trophic levels in the mesopelagic zone, where non-migratory fishes and invertebrates rely on the influx of surface-derived biomass.^[59] For instance, Antarctic krill (Euphausia superba), which exhibit pronounced DVM, serve as a primary prey for baleen whales like humpback and fin whales, channeling epipelagic energy to these large predators and sustaining their populations across polar food webs.^[60] DVM also influences dispersal and transport mechanisms within aquatic systems, promoting vertical and horizontal advection of organisms via interactions with currents. During ascent and descent, migrants can be passively transported by layered ocean currents, enhancing connectivity between distant habitats.^[61] Recent studies on larval small yellow croaker (Larimichthys polyactis) in the Yangtze River Estuary demonstrate how DVM allows juveniles to exploit faster currents at specific depths, facilitating offshore transport and recruitment success.^[13] Similarly, in the North Atlantic, copepod communities exhibit DVM-driven separation timescales that define distinct bioregions, with day-night depth partitioning altering species distributions and connectivity patterns.^[23] On a broader biogeographic scale, DVM contributes to the delineation of ocean bioregions through depth-based habitat partitioning, creating four primary zones in regions like the North Atlantic where migratory behaviors synchronize community structures diurnally.^[23] This partitioning influences species diversity gradients, as variations in migration amplitude and timing along latitudinal or depth clines modulate coexistence and resource use.^[62] Furthermore, DVM affects invasion success by imposing barriers to range expansion; for example, tropical vertical migrators face challenges crossing cold-water thermal gradients during migrations, limiting poleward invasions and shaping global distribution patterns.^[62] The ecological impacts of DVM extend to fisheries and interspecies interactions, altering predation dynamics and niche overlap. In the Barents Sea, capelin (Mallotus villosus) DVM influences cod (Gadus morhua) predation efficiency, as capelin's daytime descent into deeper waters reduces overlap with surface-oriented cod schools, affecting stock assessments and fishery yields.^[63] Partial migrations, where only subsets of populations migrate, further enable niche partitioning; in mysids (Mysis spp.), stable isotope analysis reveals that migrating individuals exploit pelagic resources while non-migrators access benthic food sources, reducing intraspecific competition and enhancing overall community stability.^[10]

Variations and anomalies

Environmental influences on patterns

Diel vertical migration (DVM) patterns vary significantly across aquatic habitats due to differences in water column depth and physical dynamics. In freshwater lakes, migration amplitudes are typically shallower, ranging from 10 to 50 meters, constrained by the limited depth and stronger thermal stratification compared to oceanic environments where amplitudes can extend from 200 to 1000 meters or more, allowing for broader vertical excursions in response to light and predation gradients.^[64]^[65] In estuarine systems, tidal cycles introduce additional synchronization that often reduces the regularity of diel patterns, as zooplankton and larval migrations align more closely with tidal flows for retention or transport, overriding pure solar-driven rhythms.^[66]^[47] Climate change exerts profound influences on DVM through ocean warming and acidification, altering migration amplitudes and cue perception. Warming waters may reduce migration amplitudes in zooplankton due to changes in temperature tolerance, as observed in freshwater systems where elevated temperatures decrease migratory range.^[26] Ocean acidification causes copepods to avoid low-pH water, potentially restricting them to shallower areas and affecting predator avoidance behaviors.^[26] Anthropogenic pollution, particularly artificial light at night, further modifies DVM timing in coastal and urban-adjacent waters. Light pollution from urban sources can alter the timing and amplitude of migrations in taxa like Daphnia by interfering with natural photoperiod signals.^[67]^[68] Recent observations highlight ongoing environmental shifts in DVM patterns. In subarctic regions, capelin (Mallotus villosus) primarily occupy 0–60 m depths both day and night, with a small fraction of juveniles exhibiting reverse DVM.^[63] Similarly, mysid shrimp demonstrate partial DVM for niche partitioning, where subsets of populations remain in benthic or pelagic zones day and night, influenced by temperature gradients and resource competition in lakes like Superior.^[10]

Unusual migrations in non-aquatic systems

While diel vertical migration (DVM) is predominantly studied in aquatic environments, analogous behaviors have been observed in non-aquatic systems, particularly among microbes and terrestrial invertebrates. In alpine snowpacks, motile microbes such as bacteria and algae exhibit DVM over distances of 10-20 cm daily, driven by phototaxis to avoid intense daytime solar radiation and diurnal meltwater flows that position them near the surface (0-3 cm) at night and deeper layers (10-20 cm) during the day. A 2025 study in a northern Japanese alpine forest documented this pattern in snow algae (e.g., Chloromonas sp.) and associated bacteria, where surface positioning at night supports nutrient access in melt layers and enhances photosynthetic activity under reduced light.^[69] Terrestrial soil invertebrates, such as collembolans (springtails), display diel shifts in vertical distribution within soil profiles, though these are not true DVM but rather responses to surface conditions. In arable soils, Arthropleona collembolans like Lepidocyrtus cyaneus show peak abundances at the soil surface from midday to midnight, correlated with higher soil-surface temperatures, with reduced surface activity (likely deeper positioning) during cooler night hours, minimizing exposure risks.^[70] In aquatic contexts, unusual DVM variants occur beyond typical zooplankton, including reverse patterns in fish. A 2025 investigation in a subarctic Greenland epipelagic ecosystem revealed that juvenile capelin (Mallotus villosus) exhibit reverse DVM, with smaller individuals (<10 cm) migrating from shallow depths (0-60 m) at night to deeper layers (80-120 m) during the day, potentially to evade cod (Gadus morhua) predation while foraging on copepods; this contrasts with the standard ascent of larger capelin.^[63] In hypoxic zones, DVM amplitudes reach extremes, as seen in the Red Sea's mesopelagic layers where scattering layers of fish and invertebrates migrate ~700 m vertically—from 400-600 m daytime depths through oxygen minima (<1.4 mL L⁻¹) to <200 m at night—tolerating low oxygen via physiological adaptations like high hemoglobin affinity.^[71] Disrupted DVM cues can lead to anomalous events, such as increased stranding risks for nearshore organisms. Artificial light pollution suppresses standard DVM in zooplankton, causing avoidance of illuminated surface layers and reduced upward migration at night, potentially heightening vulnerability to predation.^[72] Additionally, a 2025 modeling study in the Benguela Current upwelling system quantified separation timescales for vertically migrating zooplankton, showing DVM facilitates faster divergence from passive particles over seasonal timescales, potentially altering local distributions under variable currents.^[73]

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Partial diel vertical migration in an omnivorous macroinvertebrate ...
Sep 16, 2016 · In this study, we first determined whether Mysis exhibit partial DVM in Lake Champlain, USA, by sampling benthic and pelagic habitats at night.Missing: estuarine | Show results with:estuarine
[12]
(PDF) Diel vertical migration and feeding dynamics of capelin and ...
Oct 7, 2025 · This study investigates diel vertical movements and trophic interactions within a subarctic epipelagic community dominated by capelin (Mallotus ...Missing: yellow croaker
[13]
Diel Vertical Migration and Transport Pattern of Larvae and ... - MDPI
The study indicates that larvae and juveniles of the small yellow croaker change their located depth with diel vertical migration and utilize the faster speed ...Missing: subarctic capelin 2020-2025
[14]
Two hundred years of zooplankton vertical migration research
May 4, 2021 · The shorter-term diel vertical migration (DVM) has a periodicity of up to 1 day and was first described by the French naturalist Georges Cuvier ...
[15]
[PDF] Biological Oceanography : An Introduction
Because of diel vertical migrations, a comparison of day and night plankton ... Diel vertical migrations are responsible for the production of moving deep.
[16]
(PDF) Two hundred years of zooplankton vertical migration research
May 8, 2021 · The shorter‐term diel vertical migration (DVM) has a periodicity of up to 1 day and was first described by the French naturalist Georges Cuvier ...
[17]
[PDF] JOURNAL OF MARINE RESEARCH - EliScholar
Clarke-Bumpus sample pairs. That is, MOCNESS day/night sample pairs clearly documented diel migration, while Clarke-Bumpus day/night samples did not. In.
[18]
[PDF] Two hundred years of zooplankton vertical migration research
Vertical migration is a geographically and taxonomically widespread behaviour among zooplankton that spans across diel and seasonal timescales.Missing: expeditions | Show results with:expeditions
[19]
The Role of Endogenous Rhythms in Vertical Migration
May 11, 2009 · The Role of Endogenous Rhythms in Vertical Migration. Volume 43, Issue 1; John E. Harris (a1); DOI: https://doi.org/10.1017/S0025315400005324.
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Acoustic detection of zooplankton diel vertical migration behaviors ...
Apr 17, 2019 · An acoustic signal can also be obtained from an acoustic Doppler current profiler (ADCP), an instrument primarily used to study ocean ...Missing: advancements | Show results with:advancements
[21]
Zooplankton diel vertical migration in the Corsica Channel ... - OS
May 29, 2019 · The ADCP measured vertical velocity and echo intensity in the water column range between about 70 and 390 m (the bottom depth is 443 m). During ...Missing: advancements | Show results with:advancements
[22]
Expanding Tara Oceans Protocols for Underway, Ecosystemic ...
Dec 10, 2019 · Diurnal Cycles. Diel vertical migration of plankton is one of the key features to understand the fate of the primary production (Steinberg et al ...
[23]
Diel Vertical Migration Shapes North Atlantic Copepod Bioregions
Mar 24, 2025 · Aim Assessing the influence of diel vertical migration (DVM) on biogeographic patterns to improve the macroecological characterisation of ...
[24]
Global satellite-observed daily vertical migrations of ocean animals
Nov 27, 2019 · This daily excursion, referred to as diel vertical migration (DVM), is thought of primarily as an adaptation to avoid visual predators in the sunlit surface ...
[25]
Illuminating the impact of diel vertical migration on visual gene ...
Diel vertical migration (DVM) of marine animals represents one of the largest migrations on our planet. Migrating fauna are subjected to a variety of light ...Missing: technological satellite telemetry
[26]
[PDF] Impacts of Climate Change on Vertical Migration of Zooplankton ...
Aug 19, 2025 · Nearly all mesopelagic zooplankton and larvae engage in nocturnal migration, making it the most common vertical migration pattern (Bianchi ...
[27]
(PDF) Diel Patterns of Zooplankton Behavior - ResearchGate
Aug 9, 2025 · The most popular hypothesis to explain the adaptive advantage of diel vertical migration is predator avoidance, i.e., nocturnal vertical ...
[28]
[PDF] Trends in Partial Diel Vertical Migration of Mysis diluviana Along a ...
Dec 8, 2023 · Partial diel vertical migration (PDVM) means some mysids remain in the benthos at night, and others in the water column during the day. At 200m ...
[29]
[PDF] Diel Patterns of Zooplankton Behavior
In a study of vertical migration behavior of zooplankton in the English Channel,. Harris (1988) found that migrating zooplankton sometimes simply passed through.
[30]
A high-resolution modeling study on diel and seasonal vertical ...
Jan 24, 2018 · Despite diel and seasonal vertical migrations (DVM and SVM) of high-latitude zooplankton have been studied since the late-19th century, ...
[31]
https://doi.org/10.1016/j.ecolmodel.2017.12.010
[32]
Vertical distribution of Eucalanoid copepods within the Costa Rica ...
There was no evidence of daily vertical migration in the central CRD, but E. ... ontogenetic migrations (Ambler and Miller, 1987). Various studies have ...<|separator|>
[33]
https://doi.org/10.1093/plankt/fbv117
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https://doi.org/10.1111/brv.12715
[35]
Circadian Clock Involvement in Zooplankton Diel Vertical Migration
Jul 24, 2017 · We thus suggest that circadian clocks increase zooplankton fitness by optimizing the temporal trade-off between feeding and predator avoidance.
[36]
Circadian regulation of diel vertical migration (DVM) and metabolism ...
Oct 8, 2020 · Antarctic krill, Euphausia superba, (hereafter krill) are a key zooplankton species in the high-latitude ecosystem of the Southern Ocean due to ...Missing: global | Show results with:global
[37]
Genomic identification of a putative circadian system in the ...
The genome of Daphnia pulex, the only crustacean genome available for public use, provides a unique resource for identifying putative circadian proteins in this ...
[38]
[PDF] Nonlinear effects of body size and optical attenuation on Diel ...
We adopt a trait-based approach to explain Diel Vertical Migration (DVM) across a diverse assemblage of planktonic copepods, utilizing body size as a master ...Missing: scaling | Show results with:scaling
[39]
Body size‐ and season‐dependent diel vertical migration of ... - ASLO
Nov 26, 2021 · Diel vertical migration (DVM) is a common behavior performed by many zooplankton taxa in the world ocean and freshwater lakes. Larger organisms ...Acoustic Target Simulations · Results · Vertical Migration...
[40]
Diel vertical migration and endogenous swimming rhythm in ...
The results of the present experiments in total dark- ness with Asterope and Philomedes show that endogenous circadian rhythms which are as clear cut and ...
[41]
(PDF) Light and diel vertical migration: Spectral sensitivity and light ...
Aug 6, 2025 · Abstract: Ambient light levels determine the extent of diel vertical migration of many species including mysid. shrimps.<|control11|><|separator|>
[42]
A marine zooplankton community vertically structured by light across ...
Feb 24, 2021 · We selected an isolume (depth of continuous light intensity) of 10−7 µmol photons m−2 s−1, a midpoint of ranges published for the lower limit of ...
[43]
Effect of temperature on zooplankton vertical migration velocity
Upward velocities were strongly correlated with the water temperature in the migrating layer, suggesting that temperature could be a key factor controlling ...
[44]
Effects of salinity on diel vertical migration behavior in two red-tide ...
Aug 8, 2025 · We examined the effects of salinity on diel vertical migration (DVM) of two coastal flagellates, Chattonella antiqua and Karenia mikimotoi, ...<|separator|>
[45]
(PDF) Effect of the Hydrostatic Pressure on the Vertical Distribution ...
Aug 6, 2025 · An experimental study was undertaken to reveal the influence of the hydrostatic pressure on the early developmental stages of Laminaria ...
[46]
Predator specificity of kairomones in diel vertical migration of Daphnia
Aug 7, 2025 · ... kairomones in diel vertical migration of Daphnia: A chemical ... chemical cues released by predators, which initiate predator-avoidance.
[47]
Diel vertical migration and tidal influences on plankton densities in ...
Many zooplankton species perform diel vertical migration (DVM; Bandara et al ... Nocturnal or twilight migration is the most common behaviour observed among ...
[48]
Advantages from diel vertical migration can explain the dominance ...
) Light, temperature and nitrogen as interacting factors affecting diel vertical migrations of dinoflagellates in culture. J. Plankton Res. ,. 3. ,. 331. –344 ...
[49]
Vertical migration in zooplankton as a predator avoidance ... - ASLO
Abstract. Field data and laboratory feeding experiments support the hypothesis that predation can be an important factor in the adaptive significance of ...
[50]
5α-cyprinol sulfate, a bile salt from fish, induces diel vertical ... - eLife
May 2, 2019 · The finding that the amplitude of DVM increases with fish density indicates that kairomones ... kairomones in diel vertical migration of Daphnia: ...
[51]
Diel vertical migration in zooplankton: rapid individual response to ...
Abstract. While diel vertical migration in zooplankton has been shown recently to be a predator avoidance behavior, the mechanism by which predators induce.
[52]
Reverse Diel Vertical Migration: An Escape from Invertebrate ...
The marine copepod Pseudocalanus sp. exhibits an unusual reverse diel vertical migration in Dabob Bay, Washington, concurrently with a normal vertical ...<|separator|>
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Effects of Temperature on Growth of Zooplankton, and the Adaptive ...
Effects of Temperature on Growth of Zooplankton, and the Adaptive Value of Vertical Migration ... View PDF ... Citations. Cite As. Ian A. McLaren. 1963. Effects ...
[54]
A review of the adaptive significance and ecosystem consequences ...
Diel vertical migration (DVM) by zooplankton is a universal feature in all the World's oceans, as well as being common in freshwater environments.
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https://doi.org/10.1016/j.ecolmodel.2022.110003
[56]
https://doi.org/10.5194/bg-22-2181-2025
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https://doi.org/10.3389/fmars.2025.1652483
[58]
The Importance of Mesozooplankton Diel Vertical Migration for ...
Sep 12, 2019 · (2019) found that global zooplankton diel vertical migration (DVM) can increase export production by 14% annually.
[59]
The characteristics of krill swarms in relation to aggregating Antarctic ...
Nov 11, 2019 · At smaller scales, fin and humpback whales were associated with moderate and high levels of krill biomass respectively. Whales were also ...
[60]
Diel vertical migration of copepods in the tropical and subtropical ...
Copepods were always the most abundant group (86 %) and 241 species identified, although only 60 were >1 %. Smaller species predominated and Oncaea was the ...
[61]
Cold Water Serves as a Barrier to Range Expansion in Tropical ...
Nov 29, 2022 · Vertical migrators (also known as diel migrators) are species that move into deeper water (~200–400 m) during the day to hide from predators and ...
[62]
Diel vertical migration and feeding dynamics of capelin and cod in ...
Oct 4, 2025 · These patterns are shaped by a trade-off between maximizing feeding opportunities, minimizing predation risk, and managing energy expenditure.
[63]
Identifying and quantifying unexpected deep zooplankton diel ...
Dec 20, 2024 · Diel Vertical Migration (DVM), a widespread zooplankton behavior in freshwater and marine systems, affects ecological interactions and ...
[64]
Diel vertical migration of copepods and its environmental drivers in ...
Nov 2, 2020 · DVM is an important strategy for zooplankton to avoid predation risk from visually hunting predators, e.g., fish (Hays 2003; Zaret and Suffern ...Introduction · Material And Methods · DiscussionMissing: papers | Show results with:papers
[65]
Impact of tidal dynamics on diel vertical migration of zooplankton in ...
Mar 17, 2020 · This study investigates zooplankton distribution, dynamics, and factors controlling them during open-water and ice cover periods (from September 2016 to ...
[66]
Urban light pollution alters the diel vertical migration of Daphnia
Dec 1, 2017 · (2000). Urban light pollution alters the diel vertical migration of Daphnia. SIL Proceedings, 1922-2010: Vol. 27, No. 2, pp. 779-782.
[67]
The impacts of artificial light at night on the ecology of temperate and ...
Oct 30, 2023 · Use of an autonomous surface vehicle reveals small-scale diel vertical migrations of zooplankton and susceptibility to light pollution under low ...
[68]
Full article: The diel vertical migration of microbes within snowpacks ...
This study described the diel vertical migration (DVM) of microbes within a snowpack in an alpine forest in northern Japan.
[69]
Diel activity patterns in an arable collembolan community
In temperate regions, diel cycles in the activity or vertical migration of Collembola have been observed on tree trunks and shrubs (e.g. Gisin, 1943, Bauer, ...
[70]
https://www.sciencedirect.com/science/article/abs/pii/S0929139300001281
[71]
Distribution and diel vertical movements of mesopelagic scattering ...
Jun 13, 2012 · The mesopelagic zone of the Red Sea represents an extreme environment due to low food concentrations, high temperatures and low oxygen ...
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https://bmcenvsci.biomedcentral.com/articles/10.1186/s44329-025-00036-4
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Separation Timescales of Vertically Migrating Zooplankton and ...
Apr 5, 2025 · DVM by zooplankton is a highly diverse phenomenon, with migration depths and vertical migration speed varying geographically, seasonally, among ...Missing: prevalence | Show results with:prevalence<|control11|><|separator|>