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Deep scattering layer

The deep scattering layer (DSL) is a ubiquitous acoustic phenomenon in the global oceans, consisting of dense aggregations of mesopelagic organisms—primarily fish and invertebrates—that reflect and scatter sound waves, creating a distinct signature detectable by sonar and echosounders.^[1] First identified during World War II as an unexplained "false bottom" in ocean depth soundings, the DSL represents a key structural feature of the pelagic ecosystem, occupying daytime depths typically between 200 and 1000 meters before undergoing extensive vertical migrations.^[1] These layers are formed mainly by light-sensitive species such as lanternfish (myctophids) with gas-filled swim bladders, euphausiids, and other crustaceans, which contribute to the high acoustic backscatter at frequencies like 38 kHz.^[1] The organisms exhibit diel vertical migration (DVM), the largest synchronized animal movement on Earth by biomass, ascending to near-surface waters at dusk to feed on zooplankton under the cover of darkness, then descending at dawn to avoid predators and ultraviolet light exposure.^[2] This migration, driven by light penetration and predator-prey dynamics, influences ocean sound propagation, nutrient cycling, and carbon export through the biological pump, where fecal pellets and carcasses transport organic matter to deeper depths.^[1]^[2] Ecologically, DSLs serve as a critical trophic link, providing forage for higher predators including seabirds, marine mammals, and large fish, while their global prevalence—spanning from polar to tropical waters—underscores their role in sustaining biodiversity and energy transfer in the open ocean's "twilight zone."^[1] Detection relies on active acoustics, with ship-based multifrequency sonars revealing layer thickness, density, and behavior through echograms where warmer colors indicate stronger scattering returns from organism concentrations.^[2] Ongoing research highlights variations in DSL structure due to environmental factors like temperature, oxygen levels, and light attenuation, with recent studies (as of 2025) linking changes to deoxygenation and climate-driven redistribution, emphasizing their sensitivity to climate change.^[1]^[3]

History and Discovery

Initial Detection During World War II

During World War II, the U.S. Navy relied heavily on sonar systems for anti-submarine warfare to detect and track enemy vessels in the Pacific theater. As part of these efforts, the University of California Division of War Research (UCDWR) conducted extensive studies on underwater acoustics to improve sonar performance and understand sound propagation in the ocean.^[4] In 1942, UCDWR researchers C. F. Eyring, R. J. Christensen, and R. W. Raitt were aboard the USS Jasper near Guadalupe Island, Baja California, using echo-sounding equipment to measure ocean depths and investigate reverberation effects.^[5] The team detected unexpected strong acoustic returns that registered as a solid layer at depths of approximately 300–500 meters in waters exceeding 3,600 meters deep, creating the illusion of a "false bottom" or "phantom bottom" on sonar displays.^[5] This layer scattered sound waves intensely, mimicking the seabed and leading to erroneous depth readings that could mislead navigation and detection efforts. Initially, the phenomenon was attributed to instrumental artifacts or unknown physical features, as it did not align with known oceanographic data.^[6] In recognition of the discoverers, the layer was informally termed the "ECR layer," and its wartime observations heightened concerns about potential interferences in submarine detection, though no direct threats were identified. The first formal documentation of these findings appeared in a 1945 technical report issued by the UCDWR, marking the initial scientific acknowledgment of the anomaly despite its unresolved nature at the time.^[6] Subsequent wartime data hinted at daily depth variations in the layer, later explained by biological migrations, but confirmation awaited post-war analysis.^[5]

Post-War Research and Naming

Following World War II, research on the deep scattering layer shifted from wartime sonar applications to systematic scientific investigation, with institutions like the Scripps Institution of Oceanography leading peacetime expeditions to confirm its nature. The 1949-1950 Midpac Expedition, the first major postwar effort by Scripps, utilized sonar to map the layer across the central and eastern Pacific, revealing its consistent depth and diurnal vertical movements that aligned with biological patterns rather than geological features. Subsequent voyages, such as the 1951 Northern Holiday and 1952 Shellback expeditions, employed early midwater trawls to sample the layer, providing evidence that it consisted of aggregations of marine life rather than sediment or rock formations. These studies built on initial wartime detections but emphasized controlled observations to rule out abiotic causes.^[5] The nomenclature for the layer evolved during this period as understanding deepened. Initially termed the "ECR layer" after physicists C. F. Eyring, R. J. Christensen, and R. W. Raitt who detected it in 1942, or simply "false bottom" due to its misleading sonar returns, it was standardized as the "deep scattering layer" in the late 1940s through post-war analyses at Scripps. Zoologist Martin W. Johnson played a pivotal role, conducting round-the-clock acoustic monitoring in 1945 that demonstrated the layer's migration with light cycles, solidifying its biological basis and contributing to the adoption of the new term to reflect its acoustic properties from living scatterers. By the early 1950s, this naming became widespread in oceanographic literature, distinguishing it from earlier ad hoc descriptors.^[6]^[5] Early biological sampling efforts in the late 1940s and 1950s further validated these findings through direct collection. Researchers at Scripps used experimental net hauls from vessels like the Horizon during the 1952 Capricorn Expedition, capturing specimens from the layer's depth range and confirming dense schools of small marine organisms as the primary scatterers. These hauls, often conducted at night to track migrations, yielded samples that correlated precisely with sonar echoes, dispelling notions of a non-biological origin. Johnson's observations during these periods linked the layer's variability to ecological behaviors, such as synchronized vertical displacements.^[5] Key publications in the 1950s synthesized these advances, establishing the deep scattering layer as a hallmark of mesopelagic zone dynamics. Johnson's seminal 1948 paper detailed acoustic data from Pacific transects, arguing that the layer's movements indicated a biological sound-scattering mechanism tied to the ocean's twilight depths. Later works, including reports from the 1952 Capricorn and 1955 Eastropic expeditions, integrated trawl data with sonar profiles to describe the layer's role in broader oceanographic patterns, influencing subsequent studies on vertical distribution in the mesopelagic environment. These contributions, often published in journals like the Journal of Marine Research, provided foundational frameworks for linking acoustic phenomena to marine ecology.^[7]^[5]

Physical and Acoustic Properties

Depth Range and Vertical Migration

The deep scattering layer (DSL) typically resides at depths of 300–500 meters during the daytime within the mesopelagic zone, where it contributes to acoustic backscatter due to the aggregation of organisms.^[8] At night, portions of the layer ascend to shallower depths of 100–200 meters or even nearer the surface, reflecting the synchronized vertical movements of its constituents.^[9] This positioning is a key feature observed through acoustic surveys across various ocean basins, with variations influenced by regional oceanography.^[10] The DSL exhibits a prominent diel vertical migration (DVM) pattern, characterized by an upward migration beginning at dusk, allowing access to the nutrient-rich epipelagic waters for feeding, followed by a downward descent by dawn to deeper, darker depths.^[11] This migration serves primarily to balance foraging opportunities against predation risks, as the reduced light at depth during the day minimizes visibility to visual predators, while also avoiding harmful ultraviolet radiation exposure.^[12] Acoustic tracking reveals that these movements occur over distances of several hundred meters, with the layer's core often spanning 200–1000 meters overall but contracting or expanding based on the migration phase.^[13] Several environmental factors drive the DSL's vertical dynamics, including pressure gradients that increase by approximately 1 atmosphere every 10 meters of depth, influencing buoyancy adjustments during transit.^[14] Temperature changes across the thermocline, typically a drop of 10–20°C over 100–200 meters from the mixed layer to deeper waters, create thermal barriers that modulate migration extent and speed.^[15] Additionally, oxygen minimum zones (OMZs) at intermediate depths (often 200–800 meters) can constrain the layer's position, as many organisms avoid severely hypoxic regions, leading to compressed distributions in OMZ-influenced areas.^[14] Quantitative acoustic tracking indicates that DVM speeds for the DSL range from 1 to 10 meters per minute, with ascent rates often averaging 2–5 meters per minute and descent slightly faster, enabling completion of migrations within 4–8 hours.^[16]^[11]

Sound Scattering Mechanisms

The primary mechanism responsible for the strong sound scattering observed in the deep scattering layer (DSL) is the resonance of gas-filled swim bladders in mesopelagic fish, such as lanternfish (family Myctophidae). These swim bladders function as air bubbles that resonate effectively at acoustic frequencies between 10 and 50 kHz, overlapping with the operational range of standard echosounders (12–38 kHz).^[17] This resonance amplifies the backscattering cross-section by orders of magnitude compared to non-resonant structures, accounting for up to 90–95% of the total scattering in fish-dominated layers.^[18] Seminal theoretical models describe the swim bladder as a spherical gas cavity, where the resonance frequency f_r is approximated by f_r = \frac{1}{2\pi a} \sqrt{\frac{3\gamma P_0}{\rho}}, with a as the bladder radius, \gamma the adiabatic index, P_0 the hydrostatic pressure, and \rho the surrounding fluid density; at DSL depths (200–1000 m), this places resonance peaks within the sonar band for bladders of 1–5 cm radius.^[19] For smaller scatterers, such as zooplankton or juvenile fish where body dimensions are less than one-tenth of the acoustic wavelength (typically <1.25 cm at 38 kHz), scattering falls into the Rayleigh regime, characterized by weak, isotropic returns. In this regime, the volume backscattering strength s_v (in dB re 1 m⁻¹) is expressed as s_v = 10 \log_{10} (n \sigma_{bs}), where n is the numerical density of scatterers (individuals per cubic meter) and \sigma_{bs} is the backscattering cross-section of a single scatterer. The cross-section \sigma_{bs} scales with the sixth power of the scatterer radius and the fourth power of frequency (\sigma_{bs} \propto r^6 f^4), leading to rapid increases in scattering with higher frequencies but remaining subordinate to resonant contributions in the DSL.^[20]^[21] Non-resonant scatterers, including squid (via fluid-filled bodies and ink sacs) and siphonophores (via gelatinous structures with density contrasts), provide additional contributions through acoustic impedance mismatches with seawater, resulting in aggregate layer reflectivity without frequency-specific peaks. Squid scattering arises primarily from their mantle and pen, yielding lower \sigma_{bs} values (on the order of 10⁻⁶ to 10⁻⁴ m² at 38 kHz) compared to resonant fish, while siphonophore gas floats can occasionally resonate but generally contribute via fluid-like Rayleigh or geometric scattering. These elements enhance overall layer coherence, particularly in regions with low fish abundance.^[22]^[23] The DSL's typical thickness of 10–100 meters modulates echo intensity by integrating scatterers across the acoustic beam volume, with thicker layers (up to 200 m in some cases) yielding higher integrated backscattering but potentially diluting peak s_v values (often –60 to –40 dB re 1 m⁻¹). Organism orientation further influences returns, as the effective target strength varies with the angle between the incident sound wave and the scatterer's principal axis; for elongated fish or squid, broadside orientations can increase backscattering by 10–20 dB relative to head-on incidence, while random orientations in migrating schools average to isotropic-like responses.^[24]^[25]

Biological Composition

Dominant Organisms

The deep scattering layer (DSL) is predominantly composed of mesopelagic fishes, with lanternfish (family Myctophidae) serving as the primary contributors, exemplified by species in the genus Benthosema such as Benthosema glaciale and Benthosema pterotum. These small, silvery fishes, typically 3–15 cm in length, dominate the layer's biological makeup due to their high abundance and widespread distribution across oceanic realms. Lanternfish account for approximately 65% of the total mesopelagic fish biomass, underscoring their ecological prominence in the twilight zone.^[26]^[27] Complementing the fish component are other major taxa, including cephalopods such as enoploteuthid squids of the genus Abralia (e.g., Abralia veranyi), which inhabit the upper mesopelagic depths and contribute to the layer's density through their soft-bodied structures. Gelatinous zooplankton, notably siphonophores, form a significant portion of the non-fish biomass in some regions due to their colonial behaviors. Crustaceans, including euphausiids like Meganyctiphanes norvegica, amphipods, and occasionally decapods, also contribute, though they play a variable role in the overall composition. These groups collectively create the acoustically reflective aggregates observed in the DSL, with fish swim bladders enhancing detectability via resonance at common sonar frequencies.^[28]^[29]^[30] Adaptations among DSL organisms are finely tuned to the low-light, high-pressure mesopelagic realm. Lanternfish and similar fishes feature specialized bioluminescent photophores—bacterial- or protein-based light organs distributed across the body—for intraspecific communication, mate attraction, and prey detection in perpetual darkness. Gas-filled swim bladders provide neutral buoyancy, enabling efficient diel vertical migration (DVM) without excessive energy expenditure, while also facilitating acoustic backscattering. Counter-illumination, achieved by ventral photophores emitting light matching the intensity and spectrum of downwelling surface illumination, camouflages these organisms against silhouettes visible from below, reducing predation risk from upward-gazing predators. The DSL encompasses both migratory and non-migratory fractions: the migratory portion, chiefly myctophids and euphausiids, undertakes nightly ascents to 0–200 m for feeding, whereas the non-migratory fraction—often dominated by siphonophores and deeper-dwelling cephalopods—persists at 400–1000 m, maintaining stable scattering throughout the diel cycle.^[31]^[32]^[33]

Biomass and Distribution

The biomass of organisms within deep scattering layers (DSLs) represents one of the largest accumulations of animal biomass on Earth, dominated by mesopelagic fishes. Early estimates from the late 1970s and early 1980s, based on net sampling and limited surveys, placed the global biomass of mesopelagic fishes at 550–660 million tonnes, with lanternfish (Myctophidae) comprising the majority. Subsequent acoustic surveys, incorporating data from global research cruises around 2007 and advanced modeling, revised this figure substantially upward to approximately 10–15 billion tonnes for total mesopelagic fishes, with lanternfish alone estimated at 5–10 billion tonnes (about 65% of the total), reflecting their dominant role; recent reviews as of 2024 confirm a broader range of 2–16 billion tonnes for mesopelagic fish biomass overall.^[34]^[35] These revisions highlight how lanternfish, as the primary contributors to DSL biomass, dwarf the annual global fisheries catch by orders of magnitude. DSLs exhibit a broad vertical and horizontal distribution, occurring ubiquitously across the open ocean's mesopelagic zone (typically 200–1,000 m depth) and spanning latitudes from subtropical to subpolar regions, excluding extreme polar ice-covered areas. Horizontally, they form nearly continuous layers over vast expanses of the world's oceans, with acoustic backscatter indicating near-global coverage in epipelagic-to-mesopelagic transition zones. Local densities vary but can reach 10–100 individuals per cubic meter in aggregated patches, particularly during diel migrations or in response to environmental cues, as observed in midwater trawls and acoustic profiling.^[34] Factors influencing DSL abundance include gradients in surface ocean productivity, where biomass is markedly higher in upwelling areas due to elevated primary production fueling increased zooplankton and fish populations. For instance, acoustic models show a strong positive correlation between DSL backscatter intensity and annual net primary productivity, leading to denser layers in nutrient-rich coastal and equatorial upwelling systems. Ontogenetic migration further shapes distribution, with juveniles typically inhabiting shallower, more productive layers compared to deeper-dwelling adults, thereby concentrating biomass at different depths across life stages.^[34]^[36]

Ecological Significance

Role in the Marine Food Web

The deep scattering layer (DSL) serves as a critical trophic intermediary in the marine food web, bridging primary production in the sunlit surface ocean with higher-level consumers in the water column. Composed mainly of mesopelagic fishes, euphausiids, and other micronekton, DSL organisms perform diel vertical migrations, rising toward the surface at night to near-surface waters (typically 0–200 meters) to consume exported phytoplankton and zooplankton from the euphotic zone.^[8] This grazing activity positions DSL communities as primary consumers, efficiently linking basal autotrophic production to the broader pelagic ecosystem and sustaining biomass accumulation at intermediate trophic levels.^[8] A key functional role of the DSL involves the downward export of organic carbon, which bolsters the biological pump by sequestering atmospheric CO₂ in the deep ocean. As DSL organisms descend after feeding, they release fecal pellets, respiratory wastes, and dead biomass that sink rapidly to mesopelagic and bathypelagic depths, often below 1,000 meters in clear oceanic waters. This active flux mechanism contributes significantly to global carbon export, with studies estimating the migrant-mediated transport at 0.1–1 Gt C year⁻¹, enhancing the efficiency of carbon sequestration compared to passive particle sinking.^[1]^[37] Diel vertical migration also drives nutrient recycling, transporting essential elements like nitrogen and phosphorus upward to fuel surface productivity, particularly in nutrient-poor oligotrophic regions. Migrators assimilate or mobilize deep-water nutrients during daytime residence and release them via excretion and metabolic processes upon ascending, thereby alleviating nutrient limitations and supporting phytoplankton blooms that underpin the entire food web.^[38] Energy transfer from the DSL to apex predators occurs with an approximate 10% efficiency per trophic level, a standard ecological ratio that underscores the layer's indirect support for global fisheries and megafauna populations. The immense scale of DSL biomass, exceeding 5 Gt globally, provides the foundational energy reservoir for this cascade, enabling sustained productivity across oceanic ecosystems.^[39]^[40] DSL ecological roles are sensitive to climate change, with warming oceans and deoxygenation potentially altering migration patterns and reducing carbon export efficiency.^[1]

Interactions with Larger Predators

The deep scattering layer (DSL) serves as a critical prey resource for various marine megafauna, particularly during its nocturnal vertical migrations when organisms ascend toward the surface. Tunas, such as Atlantic bluefin tuna (Thunnus thynnus), frequently target mesopelagic fishes and squids within the DSL, with these components comprising a substantial portion of the diet in some populations during autumn migrations in the North Atlantic. Billfishes, including swordfish (Xiphias gladius), exhibit similar foraging behaviors, diving into the mesopelagic zone to exploit DSL aggregations, often aligning their movements with the layer's daily migrations from depths of around 500 m during the day to near-surface waters at night. Dolphins, notably Risso's dolphins (Grampus griseus) and dusky dolphins (Lagenorhynchus obscurus), perform targeted dives—such as rapid spin dives—to capture DSL-associated prey like squid and small fishes, particularly at night when the layer rises. Seabirds, relying on visual cues, opportunistically prey on DSL organisms during daytime when the layer remains at depth, though their access is limited compared to subsurface predators. DSL inhabitants employ several behavioral adaptations to mitigate predation risks from these larger predators. Diurnal vertical migration allows organisms to remain in deeper, darker waters during the day, evading visually oriented hunters like tunas, billfishes, and seabirds that are less effective in low-light conditions. Schooling behavior within the DSL further enhances survival, as evenly spaced groups of similar-sized animals—such as squid—can confuse approaching predators like dolphins by creating optical illusions or diluting individual risk, with individuals tightening formations (reducing inter-animal spacing by up to 50%) in response to threats. Bioluminescence serves as another key evasion tactic, enabling counter-illumination to blend with downwelling light or startling bursts to disorient attackers, particularly effective against fast-striking predators in the dim mesopelagic environment. The reliance of commercially valuable species on DSL prey has significant implications for fisheries management. Mesopelagic fishes and cephalopods from the DSL form a substantial portion of the diet for tunas, supporting stocks that contribute billions to global economies; for instance, in bluefin tuna, DSL-derived prey can account for a majority of caloric intake, underscoring the layer's role as a foundational forage base. This dependency highlights potential vulnerabilities, as overexploitation of surface predators could indirectly impact DSL-dependent populations through trophic cascades. Acoustic tagging studies provide direct evidence of these interactions, revealing synchronization between predator dives and DSL migrations. In the North Atlantic, tags deployed on 344 individuals across 12 pelagic species, including tunas and billfishes, showed that foraging depths often matched DSL positions, with swordfish and shortfin mako sharks (Isurus oxyrinchus) descending to 300–500 m during the day to intercept the layer and ascending nocturnally in tandem with prey. Similarly, archival tags on yellowfin tuna (Thunnus albacares) in the eastern Pacific documented vertical excursions aligned with DSL acoustics, confirming active pursuit of migrating schools for efficient foraging.

Research Methods

Acoustic Survey Techniques

Multibeam echosounders, operating across frequencies from 18 to 200 kHz, enable detailed three-dimensional mapping of deep scattering layers (DSLs) by emitting multiple narrow beams in a fan-shaped swath to capture volumetric backscatter data over wide areas. These systems, such as the Simrad EK80, utilize transducers at specific bands like 18 kHz for deep penetration and higher frequencies (e.g., 70-120 kHz) for finer resolution of layer structure, allowing real-time visualization of DSL extent, thickness, and internal heterogeneity up to depths of 1800 m.^[41]^[22] This capability supports comprehensive surveys of DSL distribution and dynamics, with echograms revealing vertical migrations as layers ascend toward the surface at night and descend during the day.^[41] Acoustic Doppler current profilers (ADCPs), typically configured at 75-150 kHz, provide complementary measurements of DSL migration velocities and biomass proxies by analyzing Doppler shifts in backscattered sound from moving scatterers. Moored or hull-mounted ADCPs record vertical velocities ranging from 10-50 mm/s during diel cycles, with descent rates often peaking 1-2 hours before sunrise and ascent rates following sunset, enabling precise tracking of layer movements over extended periods.^[42] The nautical area scattering coefficient (NASC), calculated from volume backscattering strength (s_v) using standardized sonar equations, serves as a key biomass proxy, integrating acoustic reflectivity to estimate organism density with values often exceeding 100 m²/nmi² in dense mesopelagic layers.^[42]^[43] Post-2020 integrations of artificial intelligence have enhanced real-time DSL classification by applying deep learning algorithms to multifrequency acoustic data, automating the differentiation of scatterers based on spectral signatures. Convolutional neural networks, trained on broadband target spectra from 38-200 kHz, achieve high accuracy in identifying mesopelagic fish versus non-fish components by combining empirical measurements with physics-informed simulations of scattering responses.^[44] Recent models incorporating machine learning with backscattering simulations further improve classification in data-scarce environments, processing echosounder outputs to estimate composition and abundance without extensive ground-truthing.^[45] Accurate conversion of acoustic echoes to biological metrics requires calibration via target strength (TS) models, which quantify the backscattering cross-section of individual organisms, particularly resonance effects from gas-filled swimbladders in mesopelagic fish. These models, often depth- and frequency-dependent (e.g., TS varying by -40 to -60 dB at 38 kHz), are derived from in situ broadband measurements and analytical simulations to standardize NASC interpretations across surveys.^[46] Echo interpretation in these techniques relies on fundamental sound scattering mechanisms, such as Rayleigh and resonance scattering, to link backscatter patterns to DSL composition.^[46]

Biological Sampling Approaches

Biological sampling of deep scattering layers (DSLs) primarily relies on midwater trawls and nets designed to target specific depths where acoustic backscatter indicates high organism densities. The Isaacs-Kidd midwater trawl (IKMT), a pioneering gear developed in the mid-20th century, uses a V-shaped diving vane to control depth and collect larger, active nekton such as fish and cephalopods from bathypelagic zones, often deployed at 200–1000 m to intersect DSLs.^[47] Modern iterations incorporate opening-closing mechanisms, like those in the Multiple Opening/Closing Net and Environmental Sensing System (MOCNESS), allowing precise sampling of discrete depth horizons to avoid contamination from surface or benthic layers and better capture vertically migrating organisms.^[48] These methods complement acoustic surveys by providing direct verification of the biological targets detected remotely.^[49] Remotely operated vehicles (ROVs) and submersibles have advanced DSL sampling since the 2010s, enabling in situ visual observations and targeted collections without the broad disturbance of trawling. Equipped with high-resolution cameras and manipulator arms, ROVs like those used in multi-method pelagic assessments allow for non-invasive imaging of fragile gelatinous organisms within DSLs, while suction samplers or biopsy tools collect tissue from evasive species such as squid and fish at depths up to 500 m.^[50] Submersible dives, often integrated with real-time acoustics, have documented diverse DSL communities in regions like the Arctic, confirming the presence of zooplankton and small fish that evade nets.^[28] Despite these tools, sampling DSLs presents significant challenges due to the mobility and fragility of resident organisms. Mobile species, including lanternfish and euphausiids, frequently escape trawls through mesh gaps or vertical evasion, resulting in low capture efficiencies often below 15% of the acoustic-estimated biomass.^[51] Pressure changes during ascent can cause barotrauma, disintegrating gelatinous forms or altering tissue integrity, which complicates taxonomic identification and physiological studies.^[52] Success rates remain limited, with trawls capturing less than 10% of targeted biomass in some mesopelagic assessments, underscoring the need for integrated approaches.^[53] Recent innovations in the 2020s, such as DNA metabarcoding of net-collected samples, have improved species detection by analyzing environmental DNA (eDNA) from water or bulk plankton tows, revealing uncaptured diversity in DSLs. This molecular technique identifies fish, invertebrates, and microbes across depth strata, with studies in the open ocean showing it detects up to twice as many taxa as traditional morphology-based methods from the same samples.^[54] Applied to MOCNESS tows at DSL depths, eDNA metabarcoding has elucidated vertical stratification and rare species contributions, enhancing understanding of community composition without relying solely on viable specimens.^[55]

Global and Temporal Variations

Regional Differences Across Oceans

In the Atlantic Ocean, deep scattering layers (DSLs) typically form at depths of 400-600 meters during the day, characterized by high densities of mesopelagic fishes, particularly lanternfish (Myctophidae), which contribute significantly to the layer's acoustic backscatter due to their swarming behavior and gas-filled swim bladders.^[56] This depth range reflects adaptations to the region's thermocline structure and oxygen levels, with lanternfish comprising up to 65% of the mesopelagic fish biomass in these layers.^[57] Around the Iberian Peninsula, Atlantic DSLs show a mix of migrant and non-migrant components, but the Mediterranean side exhibits distinct variations, including a more prominent non-migratory DSL at 300-600 meters, where scattering correlates with higher proportions of resident organisms like euphausiids and non-migrating fish, influenced by the semi-enclosed basin's circulation and reduced exchange with open Atlantic waters.^[57] In the Pacific Ocean, DSLs in the Clarion-Clipperton Zone (CCZ) are notably shallower, occurring at 200-400 meters during the day, a distribution shaped by the expansive oxygen minimum zone (OMZ) in this eastern central region, where low dissolved oxygen levels constrain vertical habitat and compress the layer toward higher-oxygen waters above the core OMZ.^[58] This shallower positioning contrasts with deeper global averages and highlights how OMZ intensity modulates DSL depth and micronekton composition, with acoustic surveys revealing variable migration amplitudes influenced by local productivity gradients.^[58] The Arctic Ocean presents an unexpected contrast to prior assumptions of sparse mesopelagic life in polar waters; a 2022 study documented a continuous 3,170-kilometer DSL spanning the Eurasian Basin, extending from the Atlantic inflow at depths of 100-500 meters and comprising zooplankton, small fish such as lanternfish and polar cod, and armhook squid, indicating advection of Atlantic biota supports this persistent layer despite harsh conditions.^[28] This finding challenges expectations of low biomass in ice-covered regions and underscores the role of warm Atlantic water inflows in sustaining DSLs far northward. In the Indian Ocean, DSL biomass is elevated near the equator, where monsoonal upwelling enhances nutrient availability and primary productivity, supporting denser aggregations of mesopelagic organisms in layers spanning 200-540 meters within the exclusive economic zone.^[59]

Seasonal and Diurnal Patterns

The deep scattering layer (DSL) exhibits pronounced diurnal vertical migration (DVM), with organisms ascending toward the surface at dusk and descending to deeper waters at dawn, resulting in peak acoustic scattering intensities during twilight transitions. This migration typically begins with layers returning to the surface approximately 17 minutes after sunset and descending about 21 minutes before sunrise, driven primarily by light cues to avoid predation while foraging on surface phytoplankton. Migration speeds average 6.5 cm/s upward and 7.6 cm/s downward, facilitating the full cycle over depths of 200–500 m in many regions.^[60]^[11] The amplitude of DVM varies significantly with latitude, being most robust in equatorial and tropical zones where day-night light contrasts are sharp, but weakening toward the poles due to prolonged twilight and midnight sun conditions that reduce the effective darkness cue for ascent. In high-latitude environments, such as the subpolar North Atlantic during midsummer, upward migration amplitudes can diminish progressively, with scattering layers showing reduced vertical excursions of less than 100 m compared to over 300 m in lower latitudes. This latitudinal gradient in migration intensity correlates with declining speeds poleward, from around 8–10 cm/s in the tropics to 3–4 cm/s in polar regions.^[61] Seasonally, DSL depths tend to shoal in summer due to warmer surface waters and a more pronounced, shallower thermocline that compresses habitable zones above the oxygen minimum layer, with daytime depths often around 200–250 m in temperate and subtropical areas. In contrast, winter conditions promote deeper layers, typically 300–450 m, as cooler temperatures deepen the mixed layer and weaken stratification, allowing broader vertical distribution. For instance, in the northern Arabian Sea, scattering layers reach their shallowest positions (e.g., 200 m for the upper daytime layer) during the summer southwest monsoon, when surface warming elevates the thermocline, while they deepen during the spring inter-monsoon period. Phytoplankton blooms, which peak in spring and autumn in temperate zones, further modulate these patterns by enhancing surface food availability, drawing migrators shallower during bloom periods and influencing biomass peaks in the DSL.^[62]^[62]^[63] Equatorial DSLs display greater consistency in depth and migration amplitude year-round, with minimal seasonal fluctuations tied to the stable, shallow mixed layer depth (often 20–50 m) and persistent upwelling that maintains uniform productivity. In temperate regions, however, pronounced fluctuations occur, with DSL depths varying by 100–200 m between seasons in response to mixed layer deepening in winter (up to 100–200 m) and shoaling in summer, which directly alters the vertical extent of the DSL and its scattering intensity.^[64] Long-term trends indicate potential compression of diel vertical migration (DVM) amplitudes due to ocean warming and deoxygenation, which expand hypoxic zones and restrict vertical habitat, leading to shallower overall distributions and reduced migration extents in models projecting future scenarios. Recent studies as of 2025 project significant compression of mesopelagic habitats in regions like the California Current due to these changes.^[65] Ongoing research also highlights intraseasonal variations in DSL structure driven by mesoscale eddies.^[66]

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Diel vertical migration of zooplankton and micronekton on the ...
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Jul 27, 2025 · Overall, multiple factors including organism types, size, behavior and orientation induce variable scattering echo levels (Stanton et al., 1994 ...
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Mar 19, 2021 · Lanternfishes currently represent one of the dominant groups of mesopelagic fishes in terms of abundance, biomass, and diversity.
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(PDF) Upward phosphorus transport by Daphnia diel vertical migration
Aug 10, 2025 · Conceivably, DVM could also cause nutrient transport between separated water layers. We experimentally investigated the influence of DVM of ...
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Vertical nitrogen flux from the oceanic photic zone by diel migrant ...
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Feb 7, 2014 · Abstract. With a current estimate of ~1,000 million tons, mesopelagic fishes likely dominate the world total fishes biomass.
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[PDF] Isaacs-kidd midwater trawl - DTIC
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The deep scattering layer micronektonic fish faunas of the Atlantic ...
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Mesopelagic fish biomass in the southern California current ecosystem
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Comparing eDNA metabarcoding and conventional pelagic netting ...
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Vertical stratification of environmental DNA in the open ocean ...
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Deep sea animal density and size estimated using a Dual-frequency ...
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Atlantic versus Mediterranean deep scattering layers around the ...
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Mesopelagic Scattering Layer Behaviors Across the Clarion ...
The goal of this research is to build a baseline of deep scattering layer depths and DVM behaviors within and around the CCZ using opportunistic active acoustic ...
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[PDF] Proceedings of the - Eprints@CMFRI
The bioacoustic scattoring layer of the EEZ of lndia is found in depths between 200-540 m. An additional layer is also recorded at a depth of 20-100 m in ...
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Indian Ocean mixed layer depth changes under global warming
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Mid‐summer fish behavior in a high‐latitude twilight zone - ASLO
May 19, 2023 · Correspondingly, mesopelagic scattering layers holding B. glaciale became progressively weaker and upward migration amplitudes smaller along a ...
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Seasonal and annual variability of vertically migrating scattering ...
The timing of the DVM and the formation, persistence, decay and reformation of the deep scattering layers seem to be governed by light, both solar and lunar.Missing: climate warming shifts
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Seasonal patterns in the mesopelagic fish community and ... - Nature
Oct 19, 2023 · The aim of this study was to describe seasonal echo-types and the vertical distribution of the Deep Scattering Layers (DSLs) as well as to relate them with ...
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[PDF] A Global Ocean Surface Mixed Layer monthly climatology
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Anticipating fluctuations of bigeye tuna in the Pacific Ocean from ...
Dec 24, 2022 · ... impact of ocean warming and deoxygenation. The results ... Diel migration is thus closely tied to the position of the deep scattering layer ...
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Sea-ice decline could keep zooplankton deeper for longer - Nature
Aug 28, 2023 · The models project that, in about three decades, the total time spent by zooplankton in the under-ice habitat could be reduced by up to one month, depending on ...