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Calanoida
Calanoida
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Calanoida
Unidentified species of copepod in the order Calanoida.
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Copepoda
Infraclass: Neocopepoda
Superorder: Gymnoplea
Giesbrecht, 1882 [1]
Order: Calanoida
Sars, 1903
Families

See text

Calanoida is an order of copepods, a group of arthropods commonly found as zooplankton. The order includes around 46 families with about 1800 species of both marine and freshwater copepods between them.[2]

Description

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Calanoids can be distinguished from other planktonic copepods by having first antennae at least half the length of the body and biramous second antennae.[2] However, their most distinctive anatomical trait is the presence of a joint between the fifth and sixth body segments.[3] The largest specimens reach 18 millimetres (0.71 in) long, but most do not exceed 0.5–2.0 mm (0.02–0.08 in) long.[2]

Classification

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The order Calanoida contains the following families:[4]

Ecology

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Calanoid copepods are the dominant animals in the plankton in many parts of the world's oceans, making up 55–95% of plankton samples.[2] They are therefore important in many food webs, taking in energy from phytoplankton and algae and 'repackaging' it for consumption by higher trophic level predators.[2] Many commercial fish are dependent on calanoid copepods for diet in either their larval or adult forms. Baleen whales such as bowhead whales, sei whales, right whales and fin whales rely substantially on calanoid copepods as a food source.[2]

References

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from Grokipedia
Calanoida is an order of copepods within the class Copepoda, consisting of approximately 2,000 distributed across about 43 families, and representing a dominant group of small planktonic crustaceans found in marine, brackish, and freshwater habitats worldwide. These organisms are characterized by an elongate, cylindrical body shape, long antennules that often extend beyond the caudal rami, and a life cycle that progresses from nauplius larvae through five copepodid stages to adulthood, with adults typically measuring 0.7–2.0 mm in length. Calanoid copepods play a pivotal role in aquatic food webs, functioning primarily as filter-feeding omnivores that graze on , thereby regulating and facilitating nutrient cycling, while serving as a crucial prey resource for larval and juvenile , invertebrates, and other . In marine ecosystems, they dominate zooplankton biomass and diversity, with around 75% of species being oceanic and contributing to processes like carbon export in the . Freshwater calanoids, such as those in the family Diaptomidae, are similarly abundant in lakes and streams, often inhabiting open water or benthic zones. Taxonomically, Calanoida belongs to the subclass under phylum Arthropoda, with key families including Calanidae, Aetideidae, and Eucalanidae; notable species like and Acartia spp. exemplify their ecological significance in regions such as the North Atlantic and coastal waters. Their reproduction is predominantly sexual, with females either broadcasting eggs or carrying them in one or two egg sacs, and eggs hatching into free-swimming nauplii that undergo multiple molts. High persists even in extreme environments, such as the benthopelagic zones of the at depths exceeding 3,000 meters, where undescribed species continue to be discovered through advanced sampling techniques.

Taxonomy

Etymology and History

The name Calanoida derives from the Calanus, which is New Latin for an ancient Indian philosopher (). The order was formally established by Norwegian zoologist Georg Ossian Sars in 1903, as part of his comprehensive monograph An Account of the Crustacea of , volume IV, where he synthesized observations of calanoid species from Norwegian waters. This classification built directly on foundational works by Wilhelm Giesbrecht, whose detailed monographs on ic copepods, published between 1888 and 1893 as part of the Fauna und Flora des Golfes von Neapel series and related expeditions, provided systematic illustrations and keys that highlighted calanoid diversity in marine . Early taxonomic understanding emerged in the late 18th and early 19th centuries through descriptions of individual species, such as Otto Friedrich Müller's 1785 account of Temora longicornis (originally under a different generic placement) in his Entomostraca studies of Scandinavian waters, alongside contributions from contemporaries like Carl Adolph Agardh and . Initial classifications often conflated calanoids with other groups, such as cyclopoids, due to superficial similarities in body segmentation and appendage arrangements; this confusion was progressively resolved in the late 19th and early 20th centuries through focused examinations of antennule segmentation and swimming leg morphology, as emphasized in Giesbrecht's and Sars's works. Twentieth-century advancements incorporated paleontological evidence, with revisions integrating fossil records from various deposits—that extended the group's evolutionary history and supported its ancient origins in continental and marine environments. Modern molecular phylogenetic analyses, using markers like 18S rRNA and COI, have further affirmed the and boundaries of Calanoida, refining historical delineations without altering the core order established by Sars.

Classification and Phylogeny

Calanoida is an order within the class Copepoda, subclass , subphylum Crustacea, and Arthropoda. This order encompasses approximately 43 families, 250 genera, and 2,000 described , predominantly marine but also including significant freshwater representatives. The taxonomic structure is organized into several superfamilies, including Augaptiloidea, Centropagoidea, Bathypontioidea, Eucalanoidea, Spinocalanoidea, and Clausocalanoidea, with Pseudodiaptomoidea and Diaptomoidea representing key freshwater lineages. Key families within these superfamilies include the Calanidae (e.g., Calanus finmarchicus in Eucalanoidea, a prominent North Atlantic planktonic ), Euchaetidae (also in Eucalanoidea, known for predatory forms in oceanic depths), and Diaptomidae (in Diaptomoidea, dominant in freshwater ecosystems). Phylogenetic analyses indicate that Calanoida originated from benthic or benthopelagic ancestors during the to eras, with the initial colonization of the pelagic realm occurring in the mid- around 400 million years ago. Molecular , primarily from sequences of the 18S and 28S rRNA genes, strongly supports the of Calanoida and its divergence from the sister order approximately 300–400 million years ago, aligning with the radiation of early lineages into aquatic environments. Multi-gene studies incorporating mitochondrial markers like COI and Cyt b further corroborate this timeline, revealing that basal groups such as Augaptiloidea and Centropagoidea transitioned to pelagic habitats during the , while more derived superfamilies like Clausocalanoidea emerged in the Permian amid expanding ocean oxygenation. Cladistic analyses have utilized over 50 morphological characters to resolve inter-family relationships, including antennule segmentation patterns, the presence or absence of a rostrum, and features of the female genital and swimming legs. These studies identify Epacteriscidae as the to all other calanoids, with Augaptiloidea occupying a basal position within one of two major clades that also encompasses Centropagoidea and Pseudocyclopoidea. Integration of molecular data in multi-gene phylogenies has refined these relationships, confirming the of six superfamilies (Augaptiloidea, Centropagoidea, Bathypontioidea, Eucalanoidea, Spinocalanoidea, and Clausocalanoidea) and highlighting the paraphyletic nature of Megacalanoidea, thus providing a robust framework for understanding calanoid evolutionary history.

Anatomy and Physiology

External Morphology

Calanoid copepods exhibit a distinctly segmented body divided into two main regions: the prosome and the urosome, with a major articulation point between them that allows flexibility in movement. The prosome comprises the cephalosome (fused head and first thoracic segment) plus the first four thoracic segments, totaling five , while the urosome consists of the fifth thoracic segment, the genital , and four abdominal . In females, the fifth thoracic segment is fused with the genital into a double , resulting in five urosomal overall; in males, these are separate, resulting in six urosomal . This division is characteristic of the order and facilitates hydrodynamic efficiency in planktonic lifestyles. Adult calanoids typically measure 0.5 to 2.0 mm in length, though larger species like Calanus hyperboreus can reach up to 8 mm total length, showcasing significant size variation across the order. Sexual dimorphism is evident, particularly in the antennules, where males possess longer, geniculate right antennules adapted for mate detection, often exceeding half the body length. The antennules themselves are elongate, multi-segmented appendages with 20 to 28 segments, serving sensory functions. A single naupliar eye, a remnant from larval stages, is located medially on the cephalosome, providing basic phototaxis capabilities. The rostrum, an anterior cephalic projection, is present in most families but absent in some, such as certain deep-sea forms, and aids in sensory perception. The key appendages include the biramous second antennae, which are primary limbs for and feeding via rhythmic beating to generate feeding currents. Mandibles feature gnathobases with cutting edges for mastication of prey, while maxillipeds, the most prominent thoracic appendages, function as major feeding structures with elaborate setation for particle capture. The five pairs of biramous legs (P1–P5) on the vary in structure, with P1–P4 typically three-segmented on both rami and P5 often reduced or uniramous. Caudal rami at the urosome terminus bear 4 to 6 setae, including principal propulsion setae that enhance escape responses and steering.

Internal Systems

The digestive system of calanoid copepods consists of a , , and , adapted for processing diverse particles including , , and prey. The includes a chitin-lined and a naupliar region derived from larval morphology, facilitating initial and mechanical breakdown through associated musculature. The , the primary site of and absorption, features a glandular with specialized cells such as R-cells for , D-cells for intracellular , and F-cells secreting enzymes like and ; it is divided into anterior, middle, and posterior regions, with diverticula functioning similarly to a for lipid storage and nutrient processing. The , also chitin-lined, expels waste as compact fecal pellets, with a mechanism in some aiding pellet formation and expulsion. Circulation in calanoid copepods occurs via an open hemocoel system, where hemolymph bathes the organs directly, pumped by a dorsal heart located in the second or third metasomal segment. The heart, a muscular tube with one to three ostia depending on the species, connects anteriorly to an aorta and distributes hemolymph through sinuses, supporting nutrient and oxygen transport in the absence of gills—exchange occurs via the integument and hindgut. Excretion and osmoregulation are handled by paired maxillary glands in adults, consisting of an end-sac, secretory tubule, and duct opening at the maxilla base; these glands filter hemolymph and regulate ions, a critical function for species inhabiting varied salinities, such as marine forms like Calanus spp. versus freshwater diaptomids, where active ion reabsorption prevents osmotic stress. The comprises a with protocerebral, deutocerebral, and tritocerebral lobes, from which a ventral cord extends posteriorly to innervate the body, including optic lobes connected to the persistent naupliar eye—a simple, pigmented sensitive to intensities as low as 10¹¹ photons m⁻² s⁻¹ in species like Acartia tonsa. Sensory mechanisms include chemoreceptors on the antennules, aesthetascs detecting pheromones for mate and chemical cues from sources, and mechanoreceptors responding to hydrodynamic disturbances for prey detection. Rudimentary statocysts provide balance and orientation, aiding in maintaining position during suspension feeding or escape responses. Reproductive organs in calanoid copepods include gonads located in the prosome, with development initiating in the copepodid V stage. In females, paired , often fused medially into a single structure, extend paired oviducts from the to the genital , where a genital atrium houses spermathecae for storage post-mating; eggs mature in the oviducts before release as free-spawned clutches or attached masses. In males, a single unpaired testis connects via a to a seminal vesicle and sac in the genital , enabling transfer during copulation.

Reproduction and Life Cycle

Reproductive Biology

Calanoid copepods exhibit complex behaviors that facilitate and copulation between males and females. Males actively search for females using modified antennules, often the right geniculate antennule, to detect and grasp receptive females during precopulatory pairing. Chemical cues, including pheromones released by females, play a crucial role in remote mate location and triggering male pursuit, enhancing rates in dilute planktonic environments. Once grasped, males employ their modified fifth legs as claspers to secure the female, positioning her for transfer; this precopulatory phase can involve struggle and may last from seconds to minutes depending on . typically proceeds through stages of , pursuit, capture, and copulation, with sensory structures like antennular setae aiding precise orientation. Fertilization in calanoids is internal and occurs via spermatophores, gelatinous packets of sperm produced by males and attached to the female's genital . During copulation, the male's fifth legs guide the into place on the female urosome, where it releases sperm to fertilize oocytes in the oviducts; a single often suffices for multiple egg clutches. Most species engage in broadcast spawning, releasing fertilized eggs freely into the water column, though some, such as certain Pseudodiaptomus and Eurytemora species, carry eggs in sacs attached to the urosome until hatching. Egg production strategies in calanoids include two main types: subitaneous eggs, which hatch rapidly (within days) under favorable conditions to support immediate , and diapause eggs, which enter and can remain viable for months to years, enabling survival during adverse periods like winter or low availability. Clutch sizes typically range from 20 to 100 eggs per spawning event, varying by species and modulated by environmental factors such as and abundance; higher temperatures and nutrient-rich conditions generally increase clutch size and production rates. Sex determination in calanoids is predominantly genetic, resulting in dioecious populations with a typical 1:1 , though environmental factors like can influence outcomes in some , leading to biased ratios at birth.

Developmental Stages

The development of calanoid copepods proceeds through a series of distinct post-embryonic stages, beginning with free-swimming naupliar larvae and culminating in the adult form. These stages are characterized by sequential molts, each marking morphological and physiological changes that enhance locomotion, feeding, and eventual . The naupliar phase consists of six stages (N1 to N6), which are the initial larval forms emerging from eggs. These larvae measure approximately 0.1 to 0.3 in length and possess a simple with three pairs of appendages: antennules, antennae, and mandibles. Early naupliar stages (N1 to N2 or N3) rely on internal reserves for nutrition, transitioning to active feeding on and other small particles in later stages (N3 to N6). At the end of N6, a occurs via , transforming the nauplius into the first copepodite stage, with significant reorganization of appendages and body segmentation. Following naupliar development, calanoids enter the copepodite phase, comprising six stages (C1 to C6). These stages show progressive increases in body size, from about 0.3 mm in C1 to several millimeters in C6, along with greater complexity in appendage structure and segmentation, enabling more efficient swimming and predation. The first five copepodite stages (C1 to C5) are non-reproductive juveniles, focusing on growth and dispersal, while the sixth stage (C6) represents the sexually mature adult, complete with fully developed gonads. Molting occurs at the end of each stage through ecdysis, a process involving the shedding of the exoskeleton, during which individuals are particularly vulnerable to predation due to temporary immobility. The total duration of the calanoid life cycle varies with environmental conditions, typically spanning 1 to 3 months in warm temperate or tropical waters for many species, but extending up to 1 year or more in polar regions where low temperatures slow development. Growth patterns in later copepodite stages often involve accumulation, particularly in C4 to C5, which supports —a dormant phase allowing survival under unfavorable conditions such as food scarcity or overwintering. Sexual differentiation becomes evident in these later stages, with morphological differences in appendages and body proportions appearing by C4 or C5, setting the stage for adult reproductive roles.

Ecology

Habitats and Distribution

Calanoida exhibit a , inhabiting a wide array of aquatic environments across the globe. In marine ecosystems, they dominate the plankton community, with copepods (of which calanoids form the dominant group) comprising 55–95% of individuals in many plankton samples and serving as key components of zooplankton assemblages from polar to tropical regions. This widespread presence extends to freshwater systems, where families such as Diaptomidae are prevalent in lakes and rivers, though Calanoida are less common in groundwater habitats compared to other copepod orders. Overall, approximately 75% of Calanoida species are marine, with the remainder adapted to continental waters, reflecting multiple independent invasions of freshwater habitats. In marine settings, Calanoida occupy diverse vertical strata, from epipelagic zones near the surface to bathypelagic depths exceeding 1000 m, with many undertaking diel vertical migrations spanning hundreds of meters to avoid predators or access food resources. They thrive in open oceans, coastal areas, and zones where nutrient-rich waters enhance their abundance, as observed in regions like the Angola-Benguela frontal system. Some , such as those in the Calanus, are particularly abundant in the North Atlantic and waters, while others like Acartia tonsa exhibit traits, tolerating salinities from near-freshwater to fully marine conditions (0–35 ppt). ranges from near-freezing in polar seas to over 30°C in tropical waters support their broad latitudinal distribution. Freshwater Calanoida are primarily found in temperate lakes and ponds, with notable in isolated ancient lakes such as , where species like Epischura baikalensis dominate the . These populations show high to stable, oligotrophic conditions but can tolerate environmental gradients, including pH levels from 6 to 9 and dissolved oxygen concentrations above 2 mg/L. In nutrient-enriched freshwater systems, such as those influenced by riverine inputs, their densities increase, underscoring their responsiveness to abiotic factors like oxygen and pH stability.

Trophic Interactions and Behavior

Calanoid copepods primarily engage in suspension feeding, generating feeding currents through the coordinated beating of their maxillipeds, second antennae, and maxillae to draw in prey particles from the surrounding water. These currents create an antero-ventral flow field with velocities up to 5 mm/s, enabling efficient capture of small particles without significant locomotion. Their diet consists mainly of , such as diatoms (e.g., Thalassiosira spp.), and microzooplankton, with species like Temora longicornis and Acartia spp. showing selectivity for motile or non-motile prey based on size and abundance. Clearance rates for adults typically range from 10 to 100 ml per individual per day, as observed in Temora longicornis (up to 133 ml/day) and Pseudocalanus elongatus (around 25 ml/day), varying with prey type and concentration. As key prey items in marine food webs, calanoid copepods are heavily predated upon by larvae, , and chaetognaths, which exploit their abundance in planktonic communities. To evade these predators, they perform rapid escape jumps powered by powerful thrusts from their caudal rami, achieving speeds that can exceed those of predators up to 10 times their body size. These jumps are triggered by mechanosensory detection of hydrodynamic disturbances, allowing reactions within milliseconds. Some , such as certain Calanus spp., also employ chemical defenses, releasing substances that deter predators or interfere with their sensory cues. Behavioral adaptations in calanoid copepods enhance survival and reproduction, including where individuals ascend to surface waters at night for foraging on and descend during the day to avoid visual predators in the . This migration pattern is influenced by light regimes and predator distributions, optimizing energy intake while minimizing risk. Swarming occurs in species like Heterocope septentrionalis during mating periods, clustering individuals to increase encounter rates between males and females. Additionally, rheotaxis allows copepods such as Pseudodiaptomus spp. to orient against currents, facilitating retention within productive food patches and preventing downstream displacement. Interspecific interactions among calanoid copepods often involve , particularly at high population densities, where adults and late-stage copepodites prey on eggs and nauplii, as seen in Acartia tonsa cultures exceeding 1000 individuals per liter. This behavior intensifies under food limitation, potentially regulating population growth. is rare, but some species harbor parasitic apostome like Vampyrophrya pelagica, which attach externally and feed on host tissues, often leading to mortality.

Significance

Ecological Role

Calanoid copepods play a pivotal role in regulating in aquatic ecosystems by on , often consuming a substantial fraction of daily in productive systems. In temperate coastal waters, their can account for up to 53% of phytoplankton during peak periods, with calanoids specifically contributing around 22% in some cases, thereby preventing excessive algal blooms and maintaining community balance. Through this herbivory, they facilitate carbon flux to deeper waters and the via the production of fecal pellets, which sink rapidly and contribute significantly to vertical particulate organic carbon export, sometimes dominating the flux in oligotrophic oceans. As key intermediaries in trophic linkages, calanoid copepods transfer energy from primary producers to higher trophic levels, forming a critical link in marine food webs. Their biomass often peaks during spring blooms, supporting the growth of larval fish such as , where calanoids can comprise up to 70% or more of the diet in certain spawning seasons, exemplified by high proportions of Pseudocalanus and Paracalanus in larval guts. This transfer sustains populations of commercially important like and even larger predators such as whales. Calanoid copepods act as in plankton communities, influencing by shaping the structure of lower trophic levels and serving as sensitive indicators of environmental changes. As keystone herbivores in and temperate systems, they help maintain diverse plankton assemblages, while shifts in their abundance signal , with declining calanoid proportions in copepod communities often reflecting nutrient enrichment in lakes. They also respond to shifts, exhibiting poleward range expansions of warm-water species at rates up to 231 km per decade, which alters community composition and in response to ocean warming. In terms of ecosystem services, calanoid copepods contribute to recycling through excretion, releasing and other compounds that fuel growth in surface waters, particularly in stratified environments. Their enhances by reducing biomass and indirectly influences oxygen dynamics by modulating and vertical mixing in the .

Human Applications

Calanoid copepods, particularly species like Acartia tonsa and Acartia clausi, serve as vital live feeds in for larval stages of marine fish and due to their high nutritional quality and ease of culture under controlled conditions. These copepods are cultured at densities ranging from 10^4 to 10^5 individuals per liter to optimize nauplii production for feeding, with studies showing that initial adult stocking densities of 100–500 per liter, combined with optimal algal rations, yield high egg and naupliar outputs essential for operations. Their nutritional value stems from elevated levels of omega-3 polyunsaturated fatty acids, such as (EPA) and (DHA), which support larval growth and survival; for instance, contains up to 30–40% lipids by dry weight, predominantly these essential fatty acids that enhance the health benefits in farmed . In , calanoid copepods function as model organisms for ecotoxicological studies, particularly in assessing the impacts of pollutants like polycyclic aromatic hydrocarbons (PAHs). Acartia tonsa has been widely employed in short-term tests to evaluate PAH effects on and survival, revealing concentration-dependent reductions in egg production and naupliar viability at environmentally relevant exposure levels. Additionally, genomic studies using species such as Eurytemora affinis have advanced understanding of mechanisms, with full sequencing identifying genes involved in phase I, II, and III metabolism, providing a foundation for broader genetic in copepods. Calanoid copepods play a key role in environmental monitoring as bioindicators of water quality, where shifts in species composition and abundance signal conditions like hypoxia. In hypoxic zones, such as those in coastal sediments, calanoid populations decline due to reduced oxygen tolerance, with nauplii and eggs showing high mortality below 2 mg/L dissolved oxygen, enabling their use to track deoxygenation trends in aquatic systems. In paleoceanography, sediment egg banks of calanoid copepods serve as proxies for historical environmental conditions, as viable resting eggs preserved in anoxic layers can be hatched to reconstruct past zooplankton dynamics and ecosystem responses to climate variability. Economically, lipid-rich calanoid species hold potential for , given their high fat content—often exceeding 30% of dry in species like Calanus—which includes convertible triacylglycerols suitable for conversion. However, mass culturing faces significant challenges, including on eggs and nauplii, which intensifies at high densities above 200 adults per liter and can reduce overall yields by up to 50% in intensive systems.

References

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