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Dendrobranchiata

Dendrobranchiata is a suborder of decapod crustaceans within the order Decapoda, comprising primarily marine shrimps and prawns distinguished by their uniquely branched, tree-like gills (dendrobranchiate structure), a specialized open spermatheca (thelycum) in females that facilitates external fertilization, a distinct arrangement of the first three pairs of thoracic pereiopods lacking exopods, and the production of numerous small eggs that develop externally as free-swimming naupliar larvae. The suborder includes approximately 532 valid species organized into seven families across two superfamilies: Penaeoidea (Aristeidae, Benthesicymidae, Penaeidae, Sicyoniidae, and Solenoceridae) and Sergestoidea (Luciferidae and Sergestidae).^[3] These species inhabit a wide range of marine environments, from coastal shallows and estuaries to pelagic and deep-sea habitats down to abyssal depths, with some exhibiting bioluminescence for communication or defense in the deep ocean. Ecologically, Dendrobranchiata serve as key components of food webs, acting as predators of plankton and small invertebrates while forming vital prey for larger fish, seabirds, and marine mammals;^[4] economically, families like Penaeidae include highly valued commercial species such as the whiteleg shrimp (Litopenaeus vannamei) and tiger prawns, supporting global aquaculture and fisheries that produce over 5 million metric tonnes annually (as of 2024).^[5]

Taxonomy

Nomenclature

The name Dendrobranchiata derives from the Greek words dendron (tree) and branchia (gill), alluding to the branched (dendritic) structure of the gills characteristic of this group.^[6] In English-speaking regions, species of Dendrobranchiata are commonly referred to as prawns, particularly in the United Kingdom, Australia, and New Zealand, whereas the term shrimp predominates in North America; however, both names are applied ambiguously and frequently encompass members of the related suborder Caridea as well.^[7] Historically, Dendrobranchiata were included in the obsolete taxon Natantia, established by Boas in 1880 to group swimming decapods, which also encompassed Caridea and other shrimp-like forms. The suborder Dendrobranchiata was formally recognized by Bate in 1888, distinguished from Caridea primarily by the branched gill morphology and differences in spermatophore structure, such as the complex, often winged spermatophores in Dendrobranchiata versus the simpler, spherical ones in Caridea.^[8]^[9] Dendrobranchiata holds current subordinal status within the order Decapoda, with the type genus Penaeus Fabricius, 1798, and type species Penaeus monodon Fabricius, 1798 (originally described as Cancer monodon).^[10]

Phylogenetic relationships

Dendrobranchiata occupies a basal position within the order Decapoda as the sister group to Pleocyemata, a relationship initially supported by morphological analyses and later corroborated by molecular data such as 18S rRNA sequences. This division reflects key differences in reproductive and respiratory structures, with Dendrobranchiata characterized by dendrobranchiate gills—branching, leaf-like structures—contrasting with the trichobranchiate gills of Caridea and other pleocyemates, and by external egg development rather than the brooding typical of Pleocyemata.^[11] These traits underscore the early divergence of Dendrobranchiata, positioning it as a foundational lineage in decapod evolution. The monophyly of Dendrobranchiata has been robustly confirmed through phylogenomic approaches utilizing anchored hybrid enrichment of multiple loci, resolving it as a distinct suborder alongside Pleocyemata.^[12] However, within superfamilies like Penaeoidea, recent analyses have questioned the boundaries of certain families, such as Penaeidae, suggesting potential paraphyly based on mitogenomic and multi-locus data that highlight cryptic divergences among genera.^[13] Such findings indicate ongoing refinements in internal relationships, driven by comprehensive sampling of nuclear and mitochondrial genes. The fossil record of Dendrobranchiata documents its ancient origins, extending from the Late Devonian approximately 360 million years ago to the present day, with nearly 100 extinct species contributing to an understanding of early decapod diversification. Notable early fossils include representatives from the Carboniferous Mazon Creek deposits, such as those described in Schram et al. (1978), which preserve dendrobranchiate-like morphologies and illustrate the suborder's persistence through major geological transitions despite sparse pre-Mesozoic preservation. This temporal span highlights Dendrobranchiata's evolutionary stability, with Mesozoic and Cenozoic records revealing increased diversity in marine environments.

Diversity and classification

Dendrobranchiata is a suborder of the order Decapoda within the superorder Eucarida, encompassing shrimp-like decapods primarily adapted to marine environments. The suborder is divided into two superfamilies: Penaeoidea and Sergestoidea. Penaeoidea comprises five extant families—Aristeidae, Benthesicymidae, Penaeidae, Sicyoniidae, and Solenoceridae—while Sergestoidea includes two extant families: Luciferidae and Sergestidae. In addition to the seven extant families, there are extinct families such as Aciculopodidae (Devonian) and Aegeridae (Jurassic). This classification reflects the current consensus based on morphological and molecular evidence, with seven families in total.^[14] The extant diversity of Dendrobranchiata includes approximately 540 species distributed across 47 genera. Among these, the family Penaeidae is the most speciose, containing about 221 species across multiple genera, many of which are ecologically and commercially significant in coastal and shelf habitats.^[15] Other families exhibit lower diversity; for instance, Sergestidae accounts for around 90 species in six genera, predominantly pelagic forms. Recent taxonomic revisions have been influenced by molecular phylogenetic analyses, which indicate paraphyly in certain genera within Penaeidae. For example, the traditionally broad genus Penaeus has been subdivided based on genetic evidence, with species reassigned to genera such as Fenneropenaeus and Litopenaeus to better reflect evolutionary relationships. These changes stem from studies showing distinct mitochondrial and nuclear gene lineages, challenging earlier morphological-based splits into six genera. Additionally, new species descriptions have continued post-2010, particularly in Sergestidae; notable examples include Acetes maratayama sp. nov., a cryptic species from Cananéia, Brazil, identified through integrative taxonomy in 2024.^[16] The fossil record of Dendrobranchiata reveals an ancient lineage, with extinct diversity estimated at nearly 100 species across about 10 genera. This includes purely fossil families such as Aegeridae from the Jurassic, highlighting the suborder's evolutionary persistence since the Late Devonian (~360 million years ago), with Permian and Mesozoic records showing increased diversity.

Morphology

External anatomy

Dendrobranchiata possess the caridoid facies, a body plan common among advanced malacostracans, featuring an elongated, laterally compressed form divided into a cephalothorax—where the carapace fuses with the thoracic somites—and a muscular pleon or abdomen.^[8] This structure supports their primarily pelagic or benthic lifestyles, with body sizes varying significantly across the suborder, from approximately 10 mm in small sergestid species like certain Sergestes to a maximum of 336 mm total length in the giant tiger prawn Penaeus monodon.^[17]^[18] The exoskeleton is typically robust and calcified, often adorned with spines, grooves, and carinae on the carapace and pleon for protection and hydrodynamic efficiency, though it is softer in some pelagic families like Luciferidae.^[8] The head region includes stalked compound eyes with a distinct eyestalk and cornea, usually directed laterally to provide wide visual coverage in open water environments.^[8] A prominent rostrum projects anteriorly from the carapace, varying in length and armature by family; for instance, in Penaeidae, it is well-developed with 5–11 dorsal teeth or spines, as seen in Penaeus species where 7–10 dorsal spines are typical.^[8] The antennules are biramous, consisting of three-segmented peduncles bearing dorsolateral and ventrolateral flagella for chemosensory and mechanoreceptive functions, while the antennae feature a scale-like scaphocerite and a long, multi-segmented flagellum aiding in sensory perception and balance.^[8] The thorax bears eight pairs of appendages: the first three as maxillipeds and the subsequent five as pereopods, with the first three pereopods typically chelate (pincer-like) for grasping prey or substrate, though exopods may be present or absent depending on the family (e.g., absent in Sicyoniidae).^[8] A defining external feature is the dendrobranchiate gills, which are branched or tree-like with a central axis supporting secondary and tertiary lamellae, visible along the thoracic segments beneath the carapace; this contrasts with the lamellar gills of other decapod suborders and serves as a key diagnostic trait, though adults of Luciferidae lack gills entirely.^[8] The pleon comprises six somites, each with a lateral pleuron that does not overlap the preceding one—a subtle but distinguishing feature from Caridea—with the second pleuron notably non-overlapping the first.^[8] Biramous pleopods on somites 1–5 facilitate swimming, while the sixth bears uropods that, together with the telson, form a fan-like tail for rapid propulsion; the uropodal exopod is typically larger than the endopod and lacks a diaeresis, and the telson ends in a pointed tip armed with 4 pairs of lateral spines or setae.^[8] Sexual dimorphism is evident in the pleopods, particularly the endopodites, where males possess modified structures such as the petasma on the first pair for sperm transfer and an appendix masculina on the second.^[8]

Internal anatomy

The nervous system of Dendrobranchiata consists of a dorsal brain, or supraesophageal ganglion, divided into protocerebrum, deutocerebrum, and tritocerebrum, which processes sensory input from the eyes, antennules, and antennae.^[8] This brain connects to a ventral nerve cord via circumesophageal commissures, with the cord featuring fused thoracic ganglia and six segmental pleonal ganglia that coordinate locomotion and visceral functions.^[8] Statocysts, located in the basal antennular segment, serve as organs of balance, detecting gravity and angular acceleration through statoliths composed of sand grains or secreted material.^[8] The digestive system is divided into foregut, midgut, and hindgut regions, adapted for processing a wide range of particulate food. The foregut includes a short esophagus leading to the cardiac stomach, which houses a gastric mill equipped with chitinous ossicles for grinding ingested material, and a pyloric filter that regulates passage to the midgut.^[8] The midgut features a tubular intestine surrounded by the hepatopancreas, a multifunctional gland that secretes digestive enzymes, absorbs nutrients, and stores lipids and glycogen.^[8] The hindgut, or rectum, is a straight chitin-lined tube that compacts and expels waste through the anus at the telson base.^[8] Dendrobranchiata possess an open circulatory system, with hemolymph serving as the blood equivalent and hemocyanin as the primary oxygen carrier, binding oxygen efficiently in low-oxygen environments.^[19] The heart, a muscular, triangular sac located in the pericardial sinus, pumps hemolymph through three pairs of valved ostia and distributes it via anterior and posterior arteries, including the dorsal abdominal artery that supplies the pleon.^[8] Hemolymph returns to the pericardial sinus via branchiocardiac channels from the gills, facilitating nutrient distribution and waste removal across the hemocoel.^[19] Respiration occurs in the branchial chamber, where dendrobranchiate gills—characterized by a central rachis bearing curved secondary branches with dendritic tertiary lamellae—enable efficient gas exchange through a large surface area and countercurrent flow of water and hemolymph.^[20] These gills, numbering 5–8 pairs depending on the family, are primarily respiratory but also contribute to ammonia excretion via diffusion.^[20] Osmoregulation is primarily managed by paired antennal glands, which filter hemolymph in a coelomosac, reabsorb ions and water in a labyrinthine region, and excrete urine through nephropores at the antennal bases, maintaining internal ionic balance in varying salinities.^[21] The musculature is highly developed, supporting rapid movements essential for survival. In the pleon, alternating longitudinal and transverse muscles enable powerful flexion for the tail-flip escape response, where coordinated contraction propels the shrimp backward at speeds up to several body lengths per second.^[22] These muscles, detailed in studies of species like Penaeus setiferus, include dorsal and ventral groups attached to the exoskeleton and intersegmental tendons, with sarcomere lengths around 1.8–2.0 µm for efficient contraction.

Reproduction and life cycle

Reproductive biology

Dendrobranchiata exhibit gonochoristic reproduction with distinct male and female sexes. Males possess a petasma, formed by modified endopodites of the first pleopods, which serves as the primary organ for spermatophore transfer during mating. The petasma varies in structure across families, ranging from open and flexible forms in primitive groups like Aristeidae to closed and rigid configurations in more derived taxa such as Sicyoniidae.^[23]^[24] Mating typically involves paired male-female interactions without direct intromission, where the male uses the petasma to position and attach the spermatophore to the female's thelycum. This process often occurs shortly after the female's molt, when her exoskeleton is soft, facilitating secure attachment. The spermatophore, a complex packet containing sperm and accessory fluids, is transferred externally in open-thelycum species or guided into closed receptacles in others, ensuring sperm viability until spawning. Fertilization is external, with females releasing eggs into open water where they are inseminated by stored sperm; there is no brooding, and development proceeds pelagically. Spawning generally takes place in offshore waters to optimize larval dispersal.^[25]^[26]^[23] Fecundity in Dendrobranchiata is high, reflecting their r-selected life history strategy, with females producing 100,000 to over 1,000,000 eggs per spawn depending on body size and species. For instance, in the commercially important penaeid Litopenaeus vannamei, fecundity ranges from approximately 150,000 to 442,000 eggs per female. Hermaphroditism is rare within the suborder, though protandric sex change—where individuals mature first as males before transitioning to females—has been reported in certain Sergestidae species.^[27]^[28]^[29] In temperate species, reproduction is often seasonal, synchronized with environmental cues such as rising water temperatures and increasing photoperiod to align spawning with optimal conditions for larval survival. For example, many penaeid populations in the Mediterranean exhibit peak reproductive activity from late spring to autumn, when temperatures exceed 15–20°C and day lengths lengthen. This timing ensures that naupliar larvae are released during periods of enhanced primary productivity.^[30]^[31]

Larval stages and development

Dendrobranchiata exhibit anamorphic development, characterized by gradual morphological changes through multiple larval stages rather than abrupt metamorphosis. Eggs hatch directly into free-swimming nauplius larvae, which are non-feeding and rely on yolk reserves for nutrition. These naupliar stages typically number 5 to 8, featuring simple body plans with three pairs of propulsive appendages (antennules, antennae, and mandibles) and a median nauplius eye, lasting approximately 24 to 68 hours depending on species and temperature.^[8] Following the naupliar phase, larvae transition to the zoeal stage, which includes protozoea and mysis substages and marks the onset of feeding behavior. The protozoea comprises 3 substages, during which the carapace forms, compound eyes develop and become stalked, and thoracic appendages differentiate for locomotion and feeding; uropods appear in the third substage. The mysis stage follows with 2 to 5 substages, where larvae adopt a more shrimplike form, pereiopods become functional with exopods for swimming, and pleopod buds emerge, resembling miniature adults but still planktonic.^[8] The larval sequence concludes with the postlarval (PL) or decapodid stage, where pleopods fully develop for forward swimming, exopods reduce, and larvae settle to benthic habitats. Overall, development encompasses 12 to 20 stages across families, with total duration varying from 10 to 60 days influenced by temperature and salinity; for example, in Penaeus monodon, the sequence from nauplius to postlarva takes about 14 days at 28°C, including 6 naupliar (40–55 hours), 3 protozoeal (4–6 days), and 3 mysid (3–6 days) substages.^[8]^[32] Unlike Caridea, where development is often intracapsular and larvae hatch as advanced zoeae, Dendrobranchiata nauplii hatch externally and undergo extended free-living planktonic phases, increasing vulnerability to environmental stresses. Larval mortality is particularly high during these planktonic stages due to predation, with rates declining as development progresses toward settlement. Metamorphosis is gradual without distinct transformations, guided by environmental cues such as salinity gradients that prompt postlarvae to migrate from oceanic waters to estuarine nurseries.^[8]^[33]

Distribution and habitats

Geographic distribution

Dendrobranchiata species exhibit a predominantly tropical and subtropical distribution, with the majority occurring between approximately 40°N and 40°S latitudes, reflecting their preference for warmer waters. This range encompasses key regions such as the Indo-West Pacific and the Atlantic Ocean, where families like Penaeidae dominate, with notable abundances along the west coast of Africa from Mauritania to Angola and in the coastal waters of Brazil.^[8]^[34]^[35] Although largely confined to lower latitudes, some species extend into higher latitudes, demonstrating polar extensions. For instance, the deep-sea shrimp Aristeus antennatus reaches northern limits around 45°N in the Atlantic off France, while Gennadas kempi occurs as far south as 61°S in the Antarctic Ocean. These extensions highlight the suborder's adaptability to cooler boundary environments, though diversity decreases sharply poleward. Recent studies indicate potential poleward range expansions due to ocean warming, particularly in the Southern Ocean (as of 2024).^[8]^[36] The bathymetric distribution spans from shallow coastal zones, where genera like Penaeus thrive in estuarine and nearshore areas, to abyssal depths exceeding 2000 m, as seen in Benthesicymidae species collected beyond 3000 m off Taiwan. While exclusively marine, certain taxa show limited brackish incursions during early life stages. High endemism characterizes the Indo-Pacific, a biodiversity hotspot with over 300 species, particularly in the central region bounded by the Philippines, Indonesia, and New Guinea.^[8]^[37]^[38] Biogeographic patterns are shaped by oceanographic features, including major currents like the Gulf Stream, which supports eastern Atlantic populations by transporting larvae and influencing temperature gradients. Human activities have altered ranges, with invasive spread of Litopenaeus vannamei—native to the eastern Pacific—via aquaculture escapes into Asian waters like Thailand and U.S. coastal areas.^[39]^[40]^[41]

Habitat preferences

Dendrobranchiata, comprising primarily marine decapods such as penaeid shrimps, occupy diverse habitats from coastal estuaries to deep-sea benthic zones. Juveniles of many coastal species, particularly in the family Penaeidae, preferentially utilize estuarine environments including mangroves, seagrass beds, and shallow muddy areas for nursery grounds. For instance, juvenile Penaeus merguiensis (banana prawn) inhabit mangrove-lined estuaries along the northeast Australian coast, favoring turbid, shallow waters with muddy substrates over sandy or clear areas. These habitats provide protection from predators and abundant food resources, with postlarvae actively selecting such sites upon settlement.^[42] Adult Dendrobranchiata often shift to offshore marine habitats, typically on sand or mud bottoms where they burrow or rest. In Penaeidae, adults of species like the pink shrimp Penaeus duorarum are commonly found on firm substrates such as shell-sand mixtures or calcareous mud at depths less than 50 m. Deep-sea representatives, such as those in Aristeidae (e.g., Aristeus antennatus), prefer benthic habitats at depths of 300–1000 m on soft sediments, often in areas with volcanic or muddy bottoms. Pelagic families like Sergestidae and Luciferidae inhabit open oceanic waters, with species such as Hawaiian sergestids occupying mesopelagic zones during the day (450–1200 m) and migrating shallower at night (0–300 m).^[43]^[44]^[45] Salinity tolerance varies markedly across Dendrobranchiata, reflecting their habitat diversity. Coastal penaeid species are euryhaline, tolerating wide ranges from 5–35 ppt, with optimal conditions for juveniles of P. duorarum at 10–30 ppt and postlarvae surviving 0.5–43 ppt. In contrast, oceanic and deep-sea groups like Sergestidae are largely stenohaline, requiring near-full seawater salinity (34–35.2 ppt), as seen in Hawaiian sergestid assemblages. Aristeid species also thrive in high-salinity deep waters (around 38 ppt in the Mediterranean).^[43]^[45]^[44] Temperature preferences align with latitudinal and depth distributions, with coastal species favoring warmer waters. Penaeid shrimps exhibit optimal growth and survival between 20–30°C, such as 25–28°C for Litopenaeus vannamei juveniles, though they tolerate 12–38°C. Subtropical penaeids may undertake seasonal migrations to maintain these ranges. Deep-sea Aristeidae endure cooler, stable conditions of 12.8–14.1°C, while pelagic Sergestidae experience 5–26°C across vertical migrations.^[46]^[47]^[44]^[45] Ontogenetic habitat shifts are characteristic of many Dendrobranchiata, particularly Penaeidae, where larvae remain planktonic in offshore waters before postlarvae migrate into estuarine nurseries; juveniles develop in these protected shallows, and adults relocate to deeper offshore areas. These transitions, influenced by salinity gradients and substrate availability, enhance survival across life stages.^[48]

Ecology and behavior

Feeding and diet

Dendrobranchiata exhibit opportunistic omnivorous feeding habits, consuming a diverse array of food sources including detritus, algae, and small invertebrates such as polychaetes, copepods, and molluscs.^[8] For instance, in the green tiger prawn Penaeus semisulcatus, the diet comprises primarily crustaceans (46–65% depending on sex and morphotype), diatoms (17–30%), molluscs, foraminifera, and worms, reflecting a blend of animal and plant matter.^[49] Similarly, the deep-water rose shrimp Parapenaeus longirostris preys mainly on crustaceans (58.85% frequency), foraminiferans (55.95%), polychaetes (36.63%), and molluscs (32.04%), with occasional fish and echinoderms.^[50] Stable isotope analysis indicates that the diet of Penaeus chinensis is primarily derived from benthic microalgae (90%) and marine phytoplankton (10%), with bivalves forming a core prey component.^[51] These varied diets underscore their adaptability across habitats, from estuarine shallows to deep-sea environments. Feeding mechanisms vary by lifestyle and depth. Benthic species like penaeids scrape substrates using pereopods to gather detritus and small prey, as observed in Fenneropenaeus merguiensis.^[8] Pelagic sergestids, such as those in the family Sergestidae, employ filter-feeding to capture zooplankton including euphausiids, copepods, chaetognaths, ostracods, and radiolarians.^[8] Deep-sea benthesicymids and aristeids often scavenge, incorporating fish remains, euphausiids, and debris into their diet, as seen in Gennadas and Funchalia villosa (where fish exceed 50% of biomass).^[8] The gastric mill in the foregut, featuring a denticled median tooth and lateral teeth, grinds this heterogeneous food for efficient processing. Ontogenetic shifts broaden the diet over development: larvae in protozoea and mysis stages primarily consume phytoplankton and zooplankton like rotifers and Artemia, while juveniles and adults expand to include larger benthic invertebrates and detritus.^[52] Dendrobranchiata occupy trophic levels of approximately 2.5–3.5, functioning as secondary consumers in marine food webs.^[53] They contribute significantly to nutrient cycling in estuaries by processing detritus and benthic organic matter, facilitating nitrogen and phosphorus remineralization.^[54] Foraging is predominantly nocturnal, with many species burying in sediments during the day to conserve energy and avoid predators, then emerging to probe or ambush prey at night; this pattern is evident in solenocerids like Solenocera membranacea.^[8] Seasonal variations influence intensity, such as higher consumption in spring and autumn for Parapenaeus longirostris, potentially tied to prey availability and reproductive demands.^[50]

Predation and interactions

Dendrobranchiata serve as important prey for a range of marine predators, including various fish species that consume them as a predominant component of their diet in deep-water ecosystems.^[55] In coastal and estuarine habitats, they are targeted by piscivorous fish such as snappers and groupers, as well as wading birds like herons that forage in shallow waters.^[8] Marine mammals, including seals, occasionally prey on adult shrimps during foraging in shelf areas. The larval stages are especially susceptible to predation, experiencing high mortality rates due to their small size and pelagic dispersal.^[56] To counter these threats, Dendrobranchiata employ several escape behaviors. Rapid tail-flip propulsion, powered by contraction of abdominal muscles and pleopods, allows for sudden bursts of speed to evade approaching predators. Juvenile individuals often form schools, which can confuse attackers and reduce individual risk through the dilution effect. Pelagic species further utilize camouflage via body transparency, blending with the water column to avoid visual detection by fish predators. Additionally, some deep-sea species exhibit bioluminescence for defense, such as startling predators, counterillumination for camouflage, or communication.^[57]^[58] Dendrobranchiata engage in various biotic interactions that influence their survival and population dynamics. They benefit from commensal relationships with cleaner fish, such as wrasses, which remove ectoparasites and debris from their exoskeletons in reef and estuarine settings. In shared estuarine environments, they compete with Caridea shrimps for resources like food and shelter, potentially leading to niche partitioning based on substrate preferences. As hosts to parasites, they are frequently infested by bopyrid isopods, which attach to the branchial chamber and induce effects such as castration or sex reversal in males, impairing reproduction and growth.^[59] Social behavior in Dendrobranchiata is generally solitary in adults, with individuals forming loose aggregations during feeding or resting rather than complex hierarchies. Many species undertake spawning migrations offshore, moving from estuarine nurseries to deeper waters to release eggs, a pattern that synchronizes reproduction and exposes them to varying predation risks along the route.^[60] Ecologically, Dendrobranchiata play a key role as a prey base supporting fisheries and higher trophic levels in estuarine and marine food webs. Their abundance and health serve as indicators of estuarine ecosystem condition, reflecting water quality and habitat integrity due to their sensitivity to pollution and habitat alteration.^[8]

Economic and ecological importance

Fisheries and aquaculture

Dendrobranchiata species, particularly penaeid shrimps, constitute a major component of global wild capture fisheries, with annual production reaching approximately 3.0 million tonnes in 2022 according to FAO data.^[61] These fisheries are predominantly conducted using trawl nets in tropical and subtropical marine waters, targeting species within the Penaeus genus and related groups. Leading production comes from Asia, where countries like China harvest significant volumes primarily of Litopenaeus vannamei, while Fenneropenaeus chinensis production is around 31,000 tonnes annually as of 2022.^[62] Trawl operations, while efficient for penaeid shrimps, generate substantial bycatch, often estimated at 5 to 10 kilograms of non-target species per kilogram of shrimp, including juvenile fish and other marine life that are discarded.^[63] Key commercial species in capture fisheries include the whiteleg shrimp (Litopenaeus vannamei) and tiger prawn (Penaeus monodon), which together account for a large share of tropical trawl landings.^[64] These species are harvested extensively in coastal zones of the Indo-Pacific and Atlantic, supporting both artisanal and industrial fleets. Aquaculture of Dendrobranchiata has surpassed wild capture, with global production reaching 6.4 million tonnes in 2022, driven by intensive pond systems in Asia and Latin America, and projected to reach 6 million tonnes in 2025.^[61]^[65] Litopenaeus vannamei dominates, comprising about 80-90% of farmed shrimp output as of recent estimates, while Penaeus monodon remains significant in regions like Southeast Asia.^[66] These operations rely on hatchery-reared postlarvae stocked into earthen ponds, with feeding regimes using formulated feeds to achieve high densities. However, disease outbreaks, notably white spot syndrome virus (WSSV), pose major challenges, causing mortality rates up to 100% in affected ponds and necessitating biosecurity measures like specific pathogen-free stocks.^[67] The shrimp industry generates approximately USD 70 billion in annual economic value as of 2024, encompassing production, processing, and trade, making it one of the most lucrative seafood sectors.^[68] This value stems from high international demand, with exports from major producers like Ecuador, India, and Vietnam fueling employment in coastal communities. Shrimp aquaculture experienced a boom starting in the 1970s, spurred by technological advances in hatchery propagation and pond management, leading to exponential growth from less than 100,000 tonnes in 1975 to millions by the 1990s.^[69] In response to environmental concerns, regulations have emerged, including EU import bans on shrimp from sources using unsustainable practices such as mangrove destruction or illegal antibiotics since the early 2000s.^[70]

Conservation and threats

Dendrobranchiata species face multiple anthropogenic threats that contribute to population declines and habitat degradation. Overfishing, particularly through bottom trawling, is a primary concern, as it targets commercially valuable penaeid shrimps and disrupts benthic communities.^[71] Habitat loss is another major issue, with global mangrove forests—critical nursery grounds for many penaeid larvae—experiencing approximately a 22% decline from 1985 to 2020 due to coastal development, aquaculture expansion, and conversion to agriculture, though the rate of loss has slowed in recent decades.^[72] Pollution from agricultural runoff, industrial effluents, and plastic debris further exacerbates these pressures, accumulating in shrimp tissues and impairing reproduction and growth.^[73] Climate change compounds these threats, as ocean acidification reduces larval survival and delays development in species like Pandalus borealis by altering calcification and metabolic processes.^[74] Few Dendrobranchiata species have been formally assessed by the IUCN Red List, with most categorized as Not Evaluated due to limited data on population trends.^[75] For instance, the deep-sea blue and red shrimp Aristeus antennatus, heavily exploited by trawling in the Mediterranean, shows signs of vulnerability from overexploitation, though it lacks a global IUCN status.^[76] Escaped strains from aquaculture operations, such as those of Penaeus vannamei, can hybridize with wild populations or introduce diseases, potentially outcompeting native stocks in coastal areas.^[77] Conservation efforts for Dendrobranchiata include the establishment of marine protected areas (MPAs), which help safeguard nursery habitats and reduce bycatch. In the Gulf of California, MPAs in the Upper Gulf region support populations of Penaeus californiensis by limiting trawling and protecting estuarine zones essential for juvenile growth.^[78] Sustainable certification programs, such as the Marine Stewardship Council (MSC), have been applied to several shrimp fisheries, promoting best practices like gear restrictions and stock monitoring to ensure long-term viability.^[79] Restocking initiatives using hatchery-reared juveniles have been implemented in regions like the western Pacific to bolster depleted penaeid stocks, though their ecological impacts require ongoing evaluation.^[77] Significant knowledge gaps persist, particularly for deep-sea Dendrobranchiata species, which remain understudied due to sampling challenges in remote habitats.^[80] Recent stock declines highlight these vulnerabilities; for example, catch per unit effort for penaeid shrimps in the Indian Ocean dropped amid rising fishing pressure from 2010 to 2020, indicating biomass reductions of up to 20% in some areas.^[81] As keystone species in coastal and benthic food webs, Dendrobranchiata serve as indicators of ecosystem health, linking primary producers to higher trophic levels and supporting biodiversity in mangrove and shelf environments.^[56]

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[PDF] marine biology
This species, named Penaeus monodon by Fabricius, is the type for both the genus and the family. In 1815 Rafinesque recognized that penaeid prawns were a ...<|control11|><|separator|>
[10]
Dendrobranchiata | Evolution and Phylogeny of Pancrustacea
The suborder Dendrobranchiata comprises some 500 species that all shed a non-feeding, yolk-laden nauplius. They feature a gastric mill built around a large ...
[11]
A phylogenomic framework, evolutionary timeline and genomic ...
Apr 24, 2019 · We tentatively support Dendrobranchiata and Pleocyemata, similar to transcriptomic results [22]. The polarity of the major characters separating ...
[12]
Phylogeny of penaeoid shrimps (Decapoda: Penaeoidea) inferred ...
Natural classification of Dendrobranchiata, with a key to recent genera. In: Schram, F.R. (Ed.), Crustacean Issues I. Crustacean Phylogeny. A.A. Balkema, ...
[13]
Model-based multi-locus estimation of decapod phylogeny and ...
In the present study, the phylogenetic relationships among the decapod basal suborder Dendrobranchiata and all of the currently recognized decapod infraorders ...
[14]
World Register of Marine Species - Penaeidae Rafinesque, 1815
Apr 29, 2025 · Penaeidae Rafinesque, 1815 ; Animalia (Kingdom) ; Arthropoda (Phylum) ; Crustacea (Subphylum) ; Multicrustacea (Superclass) ; Malacostraca (Class) ...<|separator|>
[15]
The Biology of Pelagic Shrimps in the Ocean - ScienceDirect.com
The body lengths of pelagic shrimps range from 10–100 mm, however a few species grow up to 130–180 mm. The epipelagic and upper mesopelagic species are ...
[16]
[PDF] Biology and ecology (of Penaeus monodon).
The maximum total length recorded was 336 mm. (Holthuis 1980), while a mature female of 307 mm from. Madagascar was reported by Crosnier (1965) as cited by.<|control11|><|separator|>
[17]
The circulatory system of Penaeus vannamei Boone, 1931 ...
Apr 4, 2020 · The branchio-pericardial sinuses bring the hemolymph back to the pericardial sinus, from where it can reenter the heart through the ostia.
[18]
[PDF] Morphological observations on the gills of dendrobranchiate shrimps
As the choice of the name Dendrobranchiata would seem to imply, gill morphology is an important morphological character separating these two major lineages of ...Missing: reproduction | Show results with:reproduction<|control11|><|separator|>
[19]
https://onlinelibrary.wiley.com/doi/full/10.1002/jmor.21117
[20]
[PDF] Muscle development in dendrobranchiate shrimp, with comparison ...
Numerous studies have examined the larval stages of dendrobranchiate shrimp (see citations in Dall et al. 1990) but few have analyzed how the internal ...
[21]
[PDF] Sperm Transfer and Storage Structures in Penaeoid Shrimps
Presumably, the spermatophores would be pressed onto the female thelycum by the petasma during copulation. How the spermatophores might be lilted into the ...
[22]
Functional morphology of the reproductive system and sperm ...
Jul 28, 2014 · In the latter case, the spermatophores are usually deposited on or in a specific female structure, termed the thelycum, for temporary or more ...
[23]
Mating structure of the blue and red shrimp, Aristeus antennatus ...
May 10, 2019 · During mating, males place spermatophores in the females' external genitalia (thelycum) with specialized enlarged coupled and folded endopods of ...
[24]
Role of the petasma and appendices masculinae during copulation ...
It is hypothesized that the male uses the petasma, supported by the endopods of the second pleopods, to hook onto the female thelycum ... REPRODUCTIVE MORPHOLOGY ...
[25]
e/7 maturation and spawning of the penaeid prawns of the ...
Heldt (1938) found the fecundity of Penaeus kerathurus (as P. trisulcatus) to be between 800,000 and 1,300,000, while Fujinaga (1963) estimated the number of ...<|separator|>
[26]
Procedure for Maturation and Spawning of Imported shrimp Litopena
As the size of females increased from 30 g to 52 g the mean number of eggs per spawning increased from 150,000 to 442,000. ... Litopenaeus vannamei and Penaeus ...
[27]
[PDF] Coordination of Reproduction and Molt in Decapods
Generally, the frequency and amplitude of changes in temperature and day length are the proximate factors controlling the temporal patterns of reproduction, ...
[28]
(PDF) Duration and timing of reproduction in decapod crustaceans ...
Aug 6, 2025 · Specifically, the observed spawning period occurred in late summer (end of August or early September), while hatching was from December to ...
[29]
None
### Summary of Larval Stages of Penaeus monodon
[30]
[PDF] Larval-Adult Matching of Deep-Sea Shrimps from the Gulf of Mexico
Sep 23, 2021 · From the suborder Dendrobranchiata, the larvae represented two families, three genera and three species.
[31]
[PDF] Dendrobranchiata: Penaeidae
Jan 31, 2021 · monodon was the largest (mean length = 18.08 ± 0.55 cm; weight = 55.01 ± 5.87 g), followed by P. kerathurus (13.04 ± 0.43 cm; 24.73 ± 1.85 g), ...Missing: Sergestes | Show results with:Sergestes
[32]
Diversity and distribution of the marine and estuarine shrimps ...
Jun 1, 2006 · Diversity and distribution of the marine and estuarine shrimps (Dendrobranchiata, Stenopodidea and Caridea) from North and Northeast Brazil.
[33]
Fisheries, Ecosystems and Biodiversity - Sea Around Us
Key Information on Blue and red shrimp (Aristeus antennatus) ; Latitude range: Northern limit: -45°S; 45°N; southern limit: -14°S 14°N ; Bottom depth range: Min.
[34]
(PDF) Diversity and Distribution of Deep-Sea Shrimps in the Ross ...
Although decapod crustaceans are widespread in the oceans, only Natantia (shrimps) are common in the Antarctic. Because remoteness, depth and ice cover restrict ...
[35]
New records of Benthesicymus Bate, 1881 (Dendrobranchiata ...
Apr 11, 2019 · Body entirely reddish to orangish red (Fig. 1). Antennular and antennal flagella orange. Scaphocerite and bascerite pinkish white to pale white.
[36]
Biogeography of the Oceans - Oxford Academic
Some 67,000 species of crustaceans are known worldwide (Ahyong et al. 2011), which comprise more than 20% of all marine species. Copepods, with more than 16,000 ...
[37]
[PDF] Geographical distribution of pelagic decapod shrimp in the Atlantic ...
Dec 16, 2014 · Ninety-one species of pelagic decapod shrimp were identified in 938 midwater-trawl collections taken between 1963 and.
[38]
Penaeus vannamei - Marine Invasions research at SERC
Penaeus vannamei was imported for aquaculture, together with other shrimp species, between 1978 and 1991. By 1991, P. vannamei was the only species cultured.
[39]
[PDF] Abundance of introduced Pacific whiteleg shrimp Penaeus ...
Oct 18, 2021 · In Thailand, this species was introduced to replace Penaeus monodon for aquaculture in 1998, and it has represented more than. 90 percent of ...
[40]
Importance of estuarine mangroves to juvenile banana prawns
Dec 1, 2012 · Offshore catches of banana prawns, Penaeus merguiensis, are correlated with the extent of mangrove forests.
[41]
[PDF] Pink Shrimp. - Habitat Suitability Index Models - DTIC
The model applies to the postlarval and juvenile life stages and is based on four habitat variables (substrate, vegetative cover, salinity, and temperature) ...
[42]
[PDF] Deep-sea distribution, biological and ecological aspects of Aristeus ...
maximum depth distribution of the rose shrimp Aristeus antennatus in the Mediterranean Sea. ... This broad depth distribution range for this species has ...
[43]
[PDF] ecology of hawaiian sergestid shrimps (penaeidea - NOAA
This paper describes the vertical distribution and migration, population size, seasonal size-frequency distribution, and diet of 20 species of sergestid ...
[44]
Effect of temperature on growth, survival, thermal behavior, and ...
Mar 28, 2018 · The optimal growth was between 25 °C and 28 °C. Survival was 100% at 22 °C, 97% at 28 °C, and 92% at 25 °C and 31 °C.Missing: Dendrobranchiata | Show results with:Dendrobranchiata
[45]
4. design criteria required by shrimp
Temperature between 26 to 30°C are generally considered best in terms of maximum yield. That is, growth is relatively fast and survival is high.Missing: preferences | Show results with:preferences
[46]
[PDF] Crustacean Biology - the NOAA Institutional Repository
Aug 1, 2020 · These generalized habitat requirements may be complicated by changes in habitat affinity associated with differing ontogenetic stages and ...
[47]
Feeding habits of green tiger prawn Penaeus semisulcatus (De ...
The results showed that the gut diet composition of both includes diatoms, dinoflagellates, crustaceans, mollusks, foraminifera, worms, and digestive material.
[48]
(PDF) Diet composition of the deep-water rose shrimp, Parapenaeus ...
Oct 9, 2025 · Crustaceans, foraminiferans, polychaetes, and molluscs constituted the most frequent preys in the diet, with respective frequencies of 58.85, ...
[49]
Trophic ecology of Penaeus chinensis (Decapoda ...
The results confirmed that bivalves served as an important diet of P.chinensis and the prawn incorporated a mixture of benthic microalgae and marine ...
[50]
THE DIET OF PRAWNS
They will eat living animal foods like Artemia, the rotifer Brachionus and prepared foods almost immediately after hatching.Missing: Dendrobranchiata | Show results with:Dendrobranchiata
[51]
Seasonal changes in the dietary patterns and reproductive aspects ...
Nov 7, 2024 · feeding ecology Given the wide distribution of this species and its significant economic importance in the Mediterranean basin, and especially ...
[52]
[PDF] Composition, Seasonality, and Life History of Decapod Shrimps in ...
Some shrimp species also play a role in detritus decomposition and nutrient cycling. ... Despite the abundance and importance of shrimp in estuaries, there is ...
[53]
Crustacean prey in the diet of fishes from deep waters of the Eastern ...
Jan 3, 2018 · Among crustacean prey, shrimps (Dendrobranchiata/Caridea) were the predominant prey for most of the examined species. The relative abundance ...
[54]
Penaeidae) and potential competitive interactions with other species ...
Aug 9, 2025 · Based on δ 15 N values, penaeid shrimps can be considered primary consumers (Al-Maslamani et al. 2007;Abreu et al. 2017) or secondary consumers ...
[55]
A review of behavioral testing in decapod shrimp (Caridea) and ...
Feb 18, 2025 · A review of behavioral testing in decapod shrimp (Caridea) and prawns (Dendrobranchiata) with applications for welfare assessment in aquaculture.Missing: gill | Show results with:gill
[56]
The Global Diversity of Parasitic Isopods Associated with ...
Apr 25, 2012 · Parasitic isopods of Bopyroidea and Cryptoniscoidea (commonly referred to as epicarideans) are unique in using crustaceans as both intermediate and definitive ...
[57]
[PDF] Social Behaviour and Recognition in Decapod Shrimps, with ...
Abstract Common in most aquatic environments, shrimps occur in a wide range of social associations with conspecific and other species, and express various ...
[58]
https://pmc.ncbi.nlm.nih.gov/articles/PMC11844257/
[59]
[PDF] Evaluating the hatchery program of a highly exploited shrimp stock ...
Mar 3, 2022 · In China,. Fenneropenaeus chinensis is a fast-growing, short-lived shrimp species with a high commercial value. F. chinensis fishery is heavily ...<|separator|>
[60]
Bycatch & Discards of Shrimp Trawling in Arabian Gulf
In the Arabian Peninsula, a study on bycatch and discards on shrimp trawling showed high levels of bycatch and discards, with a minimum of 36% bycatch and 15% ...
[61]
[PDF] The State of World Fisheries and Aquaculture 2020
Nov 21, 2019 · Marine capture production: major species and genera. 14. 4. Capture production: FAO Major Fishing Areas. 16. 5. Inland waters capture production ...
[62]
World Aquaculture Magazine - March 2025
56 MARCH 2025 • WORLD AQUACULTURE • WWW.WAS.ORG Litopenaeus vannamei accounted for 61% of the 9.5 million tons of global shrimp production in 2022 (FAO, 2022).Missing: Penaeus monodon<|separator|>
[63]
Is economies of scale driving the development in shrimp farming ...
Jan 30, 2024 · Global production of Penaeus monodon and Litopenaeus vannamei. Source: FAO, 2022. While production of P. monodon is stagnating, farmers ...
[64]
Viral disease emergence in shrimp aquaculture: origins, impact and ...
White spot syndrome virus (WSSV) is by far the most devastating pathogen of farmed shrimp. It infects all cultured penaeids and has been responsible for much of ...White Spot Syndrome Virus · Yellow Head Virus · Taura Syndrome Virus<|separator|>
[65]
Sustainable Shrimp Farming: Global Growth and Challenges
Shrimp is the most valuable traded marine product in the world today. In 2005, farmed shrimp was a 10.6 billion industry. Today, production is growing at an ...
[66]
History of shrimp farming - Responsible Seafood Advocate
Sep 1, 2011 · Shrimp farming is a young and dynamic business with a history of rapid change in response to technological advances.Missing: EU | Show results with:EU
[67]
Food Regulatory Authorities in the EU, Japan, and the United States ...
Feb 7, 2024 · Last year, RASFF included notices for thirteen (13) shipments of shrimp refused entry into the European Union because of banned antibiotics or ...
[68]
Adverse effects of shrimp fishing need to be addressed, warns UN ...
Feb 17, 2009 · Rampant overfishing, harmful trawling practices and poor management of fishing sites are causing significant damage to seabeds and endangering ...
[69]
Mangroves Fight Climate Change, but Shrimp Farming Threatens ...
Feb 21, 2024 · Why Shrimp Farming Puts Mangroves At Risk. Since 1980, between 20 and 35 percent of all mangroves have been destroyed. There are several ...
[70]
Microplastics and heavy metals in the shrimp Penaeus californiensis ...
Oct 16, 2025 · Microplastics and heavy metals in the shrimp Penaeus californiensis from the Gulf of California: Co-occurrence and tissue distribution.Missing: conservation | Show results with:conservation
[71]
Effects of ocean acidification on early life stages of shrimp (Pandalus ...
Survival of shrimp larvae was not reduced after 5 wk of exposure to pH 7.6, but a significant delay in zoeal progression (development time) was observed.Missing: impact | Show results with:impact
[72]
Aristeus antennatus, Blue and red shrimp : fisheries - SeaLifeBase
Eastern Atlantic, Mediterranean and Western Indian Ocean: from Brazil to France, south to Natal, South Africa and east to Maldives. Length at first maturity / ...Missing: latitude | Show results with:latitude
[73]
Genetic Demography of the Blue and Red Shrimp, Aristeus antennatus
Jul 1, 2022 · antennatus exists at Palamós fishing ground (Spain, northwestern Mediterranean Sea). This plan includes meshing size regulations and a temporary ...Missing: latitude | Show results with:latitude
[74]
Evaluating the Hatchery Program of a Highly Exploited Shrimp Stock ...
Mar 2, 2022 · Hatchery programs are commonly used to enhance fishery stocks, while the efforts to minimize potential negative ecological impacts have grown ...
[75]
[PDF] Blue Shrimp, Brown Shrimp, White Shrimp, Pink Shrimp, and Seabob
To address these issues, managers have established MPAs in the Upper Gulf of California (SAGARPA 2012b,. Gerrodette 2011). While MPAs have reduced bycatch of ...<|separator|>
[76]
Shrimp or Prawn | Sustainable fish - Marine Stewardship Council
Shrimp and prawn carrying the blue MSC label are certified sustainable. MSC labelled shrimp and prawn come from fisheries that have been independently assessed.
[77]
Exploring the Biodiversity of Understudied Benthic Taxa ... - Frontiers
Jun 17, 2019 · In this study, we examine the results of rare mesophotic to deep sea surveys (one rebreather, one remotely operated submersible (ROV) + net ...
[78]
Trends in Indian Ocean marine fisheries since 1950
Jan 31, 2023 · Fisheries catches in the Indian Ocean have increased consistently since 1950, in contrast to declines observed in other oceans.