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Aquatic insect
View on WikipediaThis article includes a list of references, related reading, or external links, but its sources remain unclear because it lacks inline citations. (February 2011) |
Aquatic insects or water insects live some portion of their life cycle in the water. They feed in the same ways as other insects. Some diving insects, such as predatory diving beetles, can hunt for food underwater where land-living insects cannot compete.
Breathing
[edit]Aquatic insects must get oxygen while they are under water. Almost all animals require a source of oxygen to live. Insects draw air into their bodies through spiracles, holes found along the sides of the abdomen. These spiracles are connected to tracheal tubes where oxygen can be absorbed. All aquatic insects have become adapted to their environment with the specialization of these structures, enabling:
- Simple diffusion over a relatively thin integument
- Temporary use of an air bubble
- Extraction of oxygen from water using a plastron or blood gill
- Storage of oxygen in hemoglobin and hemocyanin molecules in hemolymph[1][2]
- Taking oxygen from surface via breathing tubes (siphons)
The nymphs of the hemimetabolous orders mayflies, dragonflies and stoneflies, and the larvae of the holometabolous orders megalopterans and caddisflies, possess tracheal gills, which are outgrowths of the body wall containing a dense network of tracheae covered by a thin cuticle through which oxygen in the water can diffuse. [3][4][5]
Some insects have densely packed hairs (setae) around the spiracles that allow air to remain near, while keeping water away from, the body. The trachea open through spiracles into this air film, allowing access to oxygen. In many such cases, when the insect dives into the water, it carries a layer of air over parts of its surface, and breathes using this trapped air bubble until it is depleted, then returns to the surface to repeat the process. Other types of insects have a plastron or physical gill that can be various combinations of hairs, scales, and undulations projecting from the cuticle, which hold a thin layer of air along the outer surface of the body. In these insects, the volume of the film is small enough, and their respiration slow enough, that diffusion from the surrounding water is enough to replenish the oxygen in the pocket of air as fast as it is used. The large proportion of nitrogen in the air dissolves in water slowly and maintains the gas volume, supporting oxygen diffusion. Insects of this type only rarely need to replenish their supply of air.[6]
Other aquatic insects can remain under water for long periods due to high concentrations of hemoglobin in their hemolymph circulating freely within their body. Hemoglobin bonds strongly to oxygen molecules. [7]
A few insects such as water scorpions and mosquito larvae have breathing tubes ("siphons") with the opening surrounded by hydrofuge hairs, allowing them to breathe without having to leave the water.
Locomotion
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Aquatic insects use different methods of locomotion in water.
Orders with aquatic or semiaquatic species
[edit]- Collembola - springtails (which are not technically insects, but are closely related)
- Ephemeroptera - mayflies
- Odonata - dragonflies and damselflies
- Plecoptera - stoneflies
- Megaloptera - alderflies, fishflies, and dobsonflies
- Neuroptera - lacewings
- Coleoptera - beetles
- Hemiptera - true bugs (water striders, giant water bugs)
- Hymenoptera - ants (e.g. Polyrhachis sokolova) and wasps (e.g. Microgaster godzilla)
- Diptera - flies
- Mecoptera - scorpionflies
- Lepidoptera - moths
- Trichoptera - caddisflies
EPT insects, an acronym for Ephemeroptera, Plecoptera and Trichoptera (mayflies, stoneflies and caddisflies), are sensitive to pollutants and are used as an indicator of water quality in streams, rivers and lakes.[8]
Marine aquatic insects
[edit]Aquatic insects live mostly in freshwater habitats, and there are very few marine insect species.[9] The only true examples of pelagic insects are the sea skaters, which belongs to the order Hemiptera, and there are a few types of insects that live in the intertidal zone, including larvae of caddisflies from the family Chathamiidae,[10] the hemipteran Aepophilus bonnairei,[11] and a few other taxa.
References
[edit]- ^ Wawrowski, A.; Matthews, P. G.; Gleixner, E.; Kiger, L.; Marden, M. C.; Hankeln, T.; Burmester, T. (2012). "Characterization of the hemoglobin of the backswimmer Anisops deanei (Hemiptera)". Insect Biochemistry and Molecular Biology. 42 (9): 603–609. Bibcode:2012IBMB...42..603W. doi:10.1016/j.ibmb.2012.04.007. PMID 22575160.
- ^ Gamboa, M. (2020). "Hemocyanin and hexamerins expression in response to hypoxia in stoneflies (Plecoptera, Insecta)". Archives of Insect Biochemistry and Physiology. 105 (3) e21743. doi:10.1002/arch.21743. PMID 32979236. S2CID 221939457.
- ^ Stanley, D.; Bedick, J (1997). "Respiration in aquatic insects". Archived from the original on 2003-12-20. Retrieved 27 December 2003.
- ^ Will, Kip (2020-10-30). Field Guide to California Insects: Second Edition. Univ of California Press. ISBN 978-0-520-96357-3.
- ^ Bionomics and Ecological Services of Megaloptera Larvae (Dobsonflies, Fishflies, Alderflies)
- ^ Thorpe, W. H. (June 2008). "Plastron respiration in aquatic insects". Biological Reviews. 25 (3): 344–390. doi:10.1111/j.1469-185X.1950.tb01590.x. PMID 24538378. S2CID 44604027.
- ^ Meyer, J.R. "Respiration in Aquatic Insects". General Entomology ENT425. NC State University. Archived from the original on 2008-07-05. Retrieved 2008-04-25.
- ^ Watershed Science Institute - USDA
- ^ Why are there so few insects at sea? Deutsche Welle, 9 July 2018.
- ^ Riek, E. F. (1977). "The marine caddisfly family Chathamiidae (Trichoptera)". Australian Journal of Entomology. 15 (4): 405–419. doi:10.1111/j.1440-6055.1976.tb01724.x. ISSN 1326-6756.
- ^ Polhemus, John T. (1976). "Shore bugs (Hemiptera: Saldidae, etc.)". In Cheng, Lanna (ed.). Marine Insects (PDF). North-Holland Publishing Co. pp. 225–262. ISBN 0-444-11213-8.
- Farb, P. (1962). The Water Dwellers [LIFE]INSECTS pg. 142.
- Meyer, J.R. (2006), "Respiration in Aquatic Insects". (Accessed 25 April 2008)
- Wigglesworth, Vincent B. Sir (1964). The life of insects. Weidenfeld & Nicolson, London
External links
[edit]- Insect stages - "Some larvae, nymphs and adult insects that live in freshwater." A UK-based web site with microscopic photos of various insects and other microorganisms as well as biological information.
Aquatic insect
View on GrokipediaDefinition and Characteristics
Definition
Aquatic insects are defined as those insects that complete at least one life stage—primarily the immature larval or nymphal stages—in aquatic environments, while adults are often terrestrial and emerge from the water to reproduce or disperse.[7] They represent approximately 10% of the roughly one million described insect species, totaling around 100,000 species, and dominate as macroinvertebrates in freshwater habitats, playing key ecological roles in nutrient cycling and food webs.[7][8] These insects are secondarily aquatic, having evolved from terrestrial ancestors, and typically require both aquatic and terrestrial phases to complete their life cycles.[7] This definition distinguishes aquatic insects from fully terrestrial species, which spend their entire life cycles on land without any submerged developmental stages.[9] It also sets them apart from amphibious or semiaquatic forms, such as certain water striders (Gerridae), which primarily exploit water surfaces or margins for feeding and oviposition but do not complete a fully submerged life stage, instead visiting water only briefly without true immersion.[9] The systematic recognition of aquatic insects emerged in early entomology during the 18th century, notably through the efforts of Carl Linnaeus, who classified numerous species with aquatic immatures in his Systema Naturae (1758), integrating observations of their life histories into broader insect taxonomy.Morphological Characteristics
Aquatic insects exhibit a range of morphological adaptations that facilitate their survival in water, primarily through modifications to body shape, integument, and appendages. Many species possess streamlined bodies to minimize hydrodynamic drag, such as the flattened, dorsoventrally compressed forms seen in heptageniid mayfly nymphs and perlid stonefly nymphs, which allow efficient navigation in fast-flowing currents.[11] Hydrophobic hairs or setae cover the exoskeleton in surface-dwelling taxa like water striders (Gerridae) and whirligig beetles (Gyrinidae), enabling buoyancy and the retention of air films for movement across water surfaces.[11] Respiratory structures are prominent morphological features, with gills serving as key adaptations for oxygen uptake. Tracheal gills, thin outgrowths of the body wall richly supplied with tracheae, are common in larval stages; for instance, mayfly (Ephemeroptera) nymphs often bear plate-like or feather-like abdominal gills that enhance gas exchange when briefly referenced in respiratory contexts.[12] Stonefly (Plecoptera) nymphs typically feature tuft-like or hair-like gills on the thorax or behind the head, such as the feathery extensions at the leg bases in many species, which support diffusion in well-oxygenated streams.[13] Dragonfly (Odonata) nymphs possess internal gills within a rectal chamber at the abdomen's end, forming a branchial basket that functions in both respiration and jet propulsion. Appendages are modified for aquatic locomotion and attachment, including flattened legs for swimming in mayflies like Heptageniidae, where lateral projections reduce drag, and hooked or sucker-like tarsi in stoneflies for clinging to substrates in turbulent flows.[11] Some Coleoptera, such as elmid riffle beetles, develop plastron structures—dense fields of rigid, hydrophobic setae that trap a stable air layer against the body, acting as a physical gill for prolonged submersion.[14] These variations across orders reflect evolutionary responses to diverse aquatic pressures, with body segmentation often retaining the typical insect tripartite form (head, thorax, abdomen) but adapted for hydrophobicity and streamlining.[4]Physiological Adaptations
Respiratory Mechanisms
Aquatic insects have evolved diverse respiratory strategies to extract oxygen from aquatic environments, where dissolved oxygen levels are typically much lower than in air, often ranging from 5 to 10 mg/L compared to approximately 300 mg/L in air.[15] These mechanisms rely primarily on diffusion across thin respiratory surfaces, supplemented by active ventilation in some species, allowing them to meet metabolic demands despite hypoxic conditions.[16] Unlike terrestrial insects that use open tracheal systems connected to spiracles, many aquatic forms have adapted closed or modified systems to prevent water entry while facilitating gas exchange.[17] Cutaneous respiration, or skin breathing, occurs through diffusion across the thin, permeable cuticle in insects with closed tracheal systems lacking spiracles, such as certain diving beetle larvae (Coleoptera: Dytiscidae).[18] Oxygen from the surrounding water diffuses directly into the hemolymph and then into internal tracheae, driven by concentration gradients, while carbon dioxide diffuses outward; this method is efficient in well-oxygenated waters but limits activity in low-oxygen conditions.[15] Gill-based respiration involves external or internal gills, such as the filamentous tracheal gills on the abdomen of caddisfly larvae (Trichoptera), where oxygen diffuses across the gill surface into densely tracheated tissues.[19] These gills are often ventilated by rhythmic undulations of the abdomen, which generate water currents to maintain diffusion gradients and enhance oxygen uptake.[20] Tracheal gills, found in groups like damselfly nymphs (Odonata: Zygoptera), consist of leaf-like appendages with a thin cuticle overlying a network of tracheae, enabling direct diffusion of dissolved oxygen from water into the tracheal system.[15] These tracheal gills are modifications of the tracheal system derived from terrestrial ancestors, and their retention, along with spiracles as terrestrial atavisms, provides evidence that aquatic larvae in orders such as Odonata, Ephemeroptera, and Plecoptera represent a secondary return to aquatic environments after the initial colonization of land by insects.[7][21] Plastron respiration utilizes a stable air layer trapped by hydrophobic hairs on the body surface, functioning as a physical gill in insects like water scavenger beetles (Coleoptera: Hydrophilidae) and some hemipterans; oxygen diffuses from the water into this air film and then into the tracheae, while carbon dioxide escapes, without needing frequent replenishment.[22] This mechanism is particularly effective in moderately hypoxic waters, as the plastron maintains a constant gas volume governed by surface tension and hair structure.[23] In low-oxygen environments, aquatic insects employ behavioral adaptations such as periodic surfacing to renew air stores in bubble-breathing species or enhanced gill ventilation through abdominal pumping, as seen in mayfly nymphs (Ephemeroptera).[16] Some species, notably chironomid midge larvae (Diptera: Chironomidae), possess hemoglobin-like proteins in their hemolymph with exceptionally high oxygen affinity, allowing storage and release of oxygen to sustain metabolism during prolonged exposure to anoxic sediments.[17] These respiratory controls not only enable survival but also influence thermal tolerance and vulnerability to environmental stressors.[24]Osmoregulation and Buoyancy
Aquatic insects, primarily inhabiting freshwater environments where they are hyperosmotic to the surrounding medium, actively regulate their internal ion and water balance to counteract passive influx of water and loss of salts across their permeable integument. The Malpighian tubules, slender excretory organs branching from the hindgut, play a central role in this process by secreting a primary urine that is iso-osmotic to the hemolymph, followed by selective reabsorption of ions and water in the hindgut to produce hypo-osmotic urine for excess water excretion. In species like mosquito larvae (e.g., Aedes aegypti), specialized rectal pads facilitate this ion recovery, enabling maintenance of hemolymph osmolality around 250-300 mOsm despite ambient freshwater osmolality near 0-5 mOsm. In rarer marine or euryhaline aquatic insects, which are hypo-osmotic to saline environments, osmoregulation shifts to hypo-osmotic regulation, emphasizing salt excretion to manage osmotic water loss and ion influx. Malpighian tubules in these taxa, such as certain brine flies (Ephydra spp.), adapt by enhancing active ion transport, often coupled with active ion transport in the hindgut, allowing survival in salinities up to 100-200 ppt.[25] Euryhaline species like the mosquito Ochlerotatus taeniorhynchus possess dedicated salt-secreting glands absent in strictly freshwater forms, which actively extrude ions to tolerate brackish conditions exceeding 35 ppt.[26] These adaptations highlight the physiological plasticity in osmoregulation, with energetic costs increasing under salinity stress, as evidenced by reduced growth rates in larvae exposed to elevated ions.[27] Buoyancy control poses unique challenges for aquatic insects navigating hydrostatic pressures, with many relying on trapped air stores to achieve neutral buoyancy without specialized organs like fish swim bladders. Diving beetles (Dytiscidae), for instance, utilize gas-filled tracheal systems and subelytral air bubbles to reduce body density, enabling prolonged submersion while foraging; these bubbles, replenished at the surface, compress under pressure but maintain sufficient lift for mid-water positioning. In phantom midge larvae (Chaoborus spp.), paired gas bladders filled with atmospheric air via tracheal connections provide adjustable buoyancy, regulated by localized pH changes in the air-sac epithelium that control gas volume through mechanochemical processes in resilin bands, allowing vertical migration to evade predators.[28] Deep-water pressures, however, compress these air stores, increasing density and limiting habitat depth for most species to shallow zones less than 10 m, where surface access for bubble renewal is feasible.Taxonomy and Diversity
Major Orders
Aquatic insects are represented across multiple insect orders, with the majority exhibiting aquatic or semiaquatic stages, especially during the larval phase. These orders encompass approximately 94,000 to 130,000 described species, accounting for roughly 9-13% of the total estimated 1 million described insect species worldwide.[29][30][31] The evolutionary history of insects dates to the Devonian period around 400 million years ago, with the oldest confirmed insect fossils being wingless forms from approximately 385 million years ago; aquatic adaptations, particularly in larval stages, emerged early, as evidenced by a 370-million-year-old larval fossil from a swamp environment.[32] Primarily, aquatic insects belong to holometabolous orders (undergoing complete metamorphosis), though some paleopterous orders (incomplete metamorphosis) also feature aquatic nymphs, reflecting multiple independent colonizations of aquatic habitats. In particular, the aquatic larvae of orders such as Ephemeroptera, Odonata, and Plecoptera represent a secondary return to water after the initial terrestrial origins of insects, retaining tracheal respiratory systems including tracheal gills as modified tracheae rather than ancient gills, and spiracles as terrestrial atavisms that prove a post-land colonization reversal.[33][21] The major orders containing aquatic or semiaquatic species include:- Ephemeroptera (mayflies): Nymphs are fully aquatic, often in running waters, with short-lived terrestrial adults; representative families include Baetidae and Heptageniidae.[34]
- Odonata (dragonflies and damselflies): Nymphs are predatory and aquatic in freshwater, using gills for respiration; key examples are from families like Aeshnidae and Coenagrionidae.[34]
- Plecoptera (stoneflies): Nymphs inhabit cool, oxygenated streams, serving as indicators of water quality; examples include Perlidae and Pteronarcyidae.[34]
- Trichoptera (caddisflies): Larvae are exclusively aquatic, often building protective cases from silk and environmental materials; prominent families are Hydropsychidae and Leptoceridae.[34]
- Megaloptera: Larvae of alderflies and dobsonflies are aquatic predators in streams and lakes, with examples from Corydalidae.[29]
- Neuroptera (some species): Certain lacewings, like spongeflies in Sisyridae, have aquatic larvae that feed on freshwater sponges.[34]
- Coleoptera (beetles): Many water beetles, such as predaceous diving beetles in Dytiscidae, have aquatic larvae and adults; whirligig beetles (Gyrinidae) are semiaquatic surface dwellers.[34]
- Diptera (true flies): Larvae of families like Chironomidae (midges) and Simuliidae (blackflies) are aquatic, often in diverse freshwater habitats; Chironomidae alone comprise a significant portion of aquatic insect biomass.[34][29]
- Hemiptera (water bugs): Semiaquatic or fully aquatic bugs, including water striders (Gerridae) on surfaces and giant water bugs (Belostomatidae) as submerged predators.[35]