(Heterotardigrada)

Rough-bodied Tardigrades

Гетеротардигради

Rough-bodied Tardigrades (class Heterotardigrada) is a class of the tardigrades (water bears) that have cephalic appendages and legs with four separate but similar digits or claws on each. 444 species have been described.

The anatomy of the reproductive system is an important defining feature in distinguishing the different groups of tardigrades. Heterotardigrades have gonoducts that open to the outside through a preanal gonopore, rather than opening into the rectum as in the Eutardigrada.

Some orders of heterotardigrades are marine, others are terrestrial, but as for all tardigrades, all are aquatic in the sense that they must be surrounded by at least a film of moisture in order to be active – though they can survive in a dormant state if the habitat dries out.

Order: Arthrotardigrada

Family: Archechiniscidae

Archechiniscus: Archechiniscus marci, Archechiniscus minutus, Archechiniscus symbalanus, Archechiniscus biscaynei, Archechiniscus bahamensis.

Family: Batillipedidae

Batillipes: Batillipes acaudatus, Batillipes algharbensis, Batillipes adriaticus, Batillipes africanus, Batillipes annulatus, Batillipes bullacaudatus, Batillipes carnonensis, Batillipes crassipes, Batillipes dicrocercus, Batillipes friaufi, Batillipes gilmartini, Batillipes lesteri, Batillipes littoralis, Batillipes longispinosus, Batillipes lusitanus, Batillipes marcelli, Batillipes mirus, Batillipes noerrevangi, Batillipes orientalis, Batillipes pennaki, Batillipes philippinensis, Batillipes phreaticus, Batillipes roscoffensis, Batillipes rotundiculus, Batillipes similis, Batillipes spinicauda, Batillipes tridentatus, Batillipes tubernatis.

Family: Coronarctidae

Coronarctus: Coronarctus disparilis, Coronarctus fastigatus, Coronarctus laubieri, Coronarctus mexicus, Coronarctus stylisetus, Coronarctus tenellus, Coronarctus verrucatus.

Trogloarctus: Trogloarctus trionyches.

Family: Halechiniscidae

Dipodarctus: Dipodarctus anaholiensis, Dipodarctus borrori, Dipodarctus subterraneus.

Clavarctus: Clavarctus falculus.

Euclavarctus: Euclavarctus convergens, Euclavarctus thieli.

Exoclavarctus: Exoclavarctus dineti.

Moebjergarctus: Moebjergarctus manganis, Moebjergarctus clarionclippertonensis.

Parmursa: Parmursa fimbriata, Parmursa torquata.

Proclavarctus: Proclavarctus fragilis.

Florarctus: Florarctus acer, Florarctus antillensis, Florarctus asper, Florarctus cervinus, Florarctus cinctus, Florarctus glareolus, Florarctus heimi, Florarctus hulingsi, Florarctus kwoni, Florarctus pulcher, Florarctus salvati, Florarctus stellatus, Florarctus vulcanius.

Ligiarctus: Ligiarctus eastwardi.

Wingstrandarctus: Wingstrandarctus corallinus, Wingstrandarctus crypticus, Wingstrandarctus intermedius, Wingstrandarctus stinae, Wingstrandarctus unsculptus.

Chrysoarctus: Chrysoarctus briandi, Chrysoarctus flabellatus.

Halechiniscus: Halechiniscus chafarinensis, Halechiniscus churakaagii, Halechiniscus greveni, Halechiniscus guiteli, Halechiniscus janus, Halechiniscus jejuensis, Halechiniscus macrocephalus, Halechiniscus paratuleari, Halechiniscus perfectus, Halechiniscus remanei, Halechiniscus tuleari, Halechiniscus yanakaagii.

Paradoxipus: Paradoxipus orzeliscoides.

Orzeliscus: Orzeliscus belopus, Orzeliscus septentrionalis, Orzeliscus asiaticus.

Opydorscus: Opydorscus fonsecae.

Angursa: Angursa abyssalis, Angursa antarctica, Angursa bicuspis, Angursa capsula, Angursa clavifera, Angursa lanceolata, Angursa lingua, Angursa seisuimaruae.

Bathyechiniscus: Bathyechiniscus tetronyx.

Lepoarctus: Lepoarctus coniferus.

Paratanarctus: Paratanarctus kristenseni.

Pleocola: Pleocola limnoriae.

Raiarctus: Raiarctus aureolatus, Raiarctus colurus, Raiarctus jesperi, Raiarctus katrinae, Raiarctus variabilis.

Rhomboarctus: Rhomboarctus aslaki, Rhomboarctus duplicicaudatus, Rhomboarctus thomassini.

Styraconyx: Styraconyx craticuliformis, Styraconyx craticulus, Styraconyx hallasi, Styraconyx haploceros, Styraconyx kristenseni, Styraconyx nanoqsunguak, Styraconyx paulae, Styraconyx qivitoq, Styraconyx robertoi, Styraconyx sardiniae, Styraconyx sargassi, Styraconyx takeshii, Styraconyx testudo, Styraconyx turbinarium, Styraconyx tyrrhenus, Styraconyx vargasi.

Tetrakentron: Tetrakentron synaptae.

Tholoarctus: Tholoarctus natans, Tholoarctus oleseni.

Actinarctus: Actinarctus doryphorus, Actinarctus lyrophorus, Actinarctus neretinus, Actinarctus physophorus.

Tanarctus: Tanarctus arborspinosus, Tanarctus breedyae, Tanarctus bubulubus, Tanarctus dendriticus, Tanarctus diplocerus, Tanarctus gracilis, Tanarctus helleouetae, Tanarctus heterodactylus, Tanarctus hirsutospinosus, Tanarctus ittanmomen, Tanarctus longisetosus, Tanarctus minotauricus, Tanarctus ramazzottii, Tanarctus tauricus, Tanarctus velatus.

Zioella: Zioella pavonina.

Family: Neoarctidae

Neoarctus: Neoarctus primigenius.

Family: Neostygarctidae

Neostygarctus: Neostygarctus acanthophorus, Neostygarctus grossmeteori, Neostygarctus lovedeluxe, Neostygarctus oceanopolis.

Family: Renaudarctidae

Renaudarctus: Renaudarctus fossorius, Renaudarctus psammocryptus.

Family: Stygarctidae

Megastygarctides: Megastygarctides christinae, Megastygarctides gerdae, Megastygarctides isounguis, Megastygarctides orbiculatus, Megastygarctides setoloso, Megastygarctides sezginii.

Parastygarctus: Parastygarctus biungulatus, Parastygarctus higginsi, Parastygarctus mediterranicus, Parastygarctus renaudae, Parastygarctus robustus, Parastygarctus sterreri, Parastygarctus svennevigi.

Pseudostygarctus: Pseudostygarctus apuliae, Pseudostygarctus galloae, Pseudostygarctus mirabilis, Pseudostygarctus rugosus, Pseudostygarctus triungulatus.

Stygarctus: Stygarctus abornatus, Stygarctus ayatori, tygarctus bradypus, Stygarctus gourbaultae, Stygarctus granulatus, Stygarctus keralensis, Stygarctus lambertii, Stygarctus spinifer.

Order: Echiniscoidea

Family: Carphanidae

Carphania: Carphania fluviatilis.

Family: Echiniscidae

Antechiniscus: Antechiniscus conversus, Antechiniscus jermani, Antechiniscus lateromamillatus, Antechiniscus moscali, Antechiniscus parvisentus, Antechiniscus perplexus.

Bryochoerus: Bryochoerus intermedius.

Bryodelphax: Bryodelphax aaseae, Bryodelphax alzirae, Bryodelphax amphoterus, Bryodelphax arenosus, Bryodelphax asiaticus, Bryodelphax atlantis, Bryodelphax australasiaticus, Bryodelphax brevidentatus, Bryodelphax crossotus, Bryodelphax decoratus, Bryodelphax dominicanus, Bryodelphax iohannis, Bryodelphax kristenseni, Bryodelphax lijiangensis, Bryodelphax maculatus, Bryodelphax mareki, Bryodelphax mateusi, Bryodelphax meronensis, Bryodelphax nigripunctatus, Bryodelphax olszanowskii, Bryodelphax ortholineatus, Bryodelphax parvulus, Bryodelphax parvuspolaris, Bryodelphax sinensis, Bryodelphax tatrensis, Bryodelphax weglarskae.

Cornechiniscus: Cornechiniscus brachycornutus, Cornechiniscus ceratophorus, Cornechiniscus cornutus, Cornechiniscus holmeni, Cornechiniscus lobatus, Cornechiniscus madagascariensis, Cornechiniscus mystacinus, Cornechiniscus schrammi, Cornechiniscus imperfectus, Cornechiniscus subcornutus, Cornechiniscus tibetanus.

Echiniscus: Echiniscus africanus, Echiniscus aliquantillus, Echiniscus angolensis, Echiniscus apuanus, Echiniscus arcangelii, Echiniscus arctomys, Echiniscus arthuri, Echiniscus aonikenk, Echiniscus azoricus, Echiniscus baius, Echiniscus baloghi, Echiniscus barbarae, Echiniscus batramiae, Echiniscus becki, Echiniscus bigranulatus, Echiniscus bisculptus, Echiniscus blumi, Echiniscus calcaratus, Echiniscus calvus, Echiniscus canadensis, Echiniscus canedoi, Echiniscus capillatus, Echiniscus carsicus, Echiniscus carusoi, Echiniscus cavagnaroi, Echiniscus cervicornis, Echiniscus charrua, Echiniscus cheonyoungi, Echiniscus cirinoi, Echiniscus clavispinosus, Echiniscus clevelandi, Echiniscus columinis, Echiniscus corrugicaudatus, Echiniscus crassispinosus, Echiniscus curiosus, Echiniscus dariae, Echiniscus darienae, Echiniscus dearmatus, Echiniscus dikenli, Echiniscus diploglyptus, Echiniscus divergens, Echiniscus dreyfusi, Echiniscus duboisi, Echiniscus egnatiae, Echiniscus ehrenbergi, Echiniscus elaeinae, Echiniscus elegans, Echiniscus evelinae, Echiniscus filamentosus, Echiniscus ganczareki, Echiniscus glaber, Echiniscus granulatus, Echiniscus heterospinosus, Echiniscus hexacanthus, Echiniscus hoonsooi, Echiniscus horningi, Echiniscus inocelatus, Echiniscus insuetus, Echiniscus jagodici, Echiniscus jamesi, Echiniscus japonicus, Echiniscus jenningsi, Echiniscus kerguelensis, Echiniscus knowltoni, Echiniscus kofordi, Echiniscus kosickii, Echiniscus lapponicus, Echiniscus laterosetosus, Echiniscus laterospinosus, Echiniscus latifasciatus, Echiniscus lichenorum, Echiniscus limai, Echiniscus lineatus, Echiniscus longispinosus, Echiniscus loxophthalmus, Echiniscus madonnae, Echiniscus maesi, Echiniscus malpighii, Echiniscus manuelae, Echiniscus marcusi, Echiniscus marginatus, Echiniscus marginoporus, Echiniscus markezi, Echiniscus marleyi, Echiniscus mauccii, Echiniscus mediantus, Echiniscus meridionalis, Echiniscus merokensis, Echiniscus migiurtinus, Echiniscus mihelcici, Echiniscus militaris, Echiniscus molluscorum, Echiniscus moniliatus, Echiniscus montanus, Echiniscus mosaicus, Echiniscus multispinosus, Echiniscus murrayi, Echiniscus nelsonae, Echiniscus nepalensis, Echiniscus nigripustulus, Echiniscus nobilis, Echiniscus oihonnae, Echiniscus ollantaytamboensis, Echiniscus osellai, Echiniscus pajstunensis, Echiniscus palmai, Echiniscus pellucidus, Echiniscus perarmatus, Echiniscus peruvianus, Echiniscus perviridis, Echiniscus phocae, Echiniscus polygonalis, Echiniscus pooensis, Echiniscus porabrus, Echiniscus postojnensis, Echiniscus pseudelegans, Echiniscus pseudowendti, Echiniscus punctus, Echiniscus pusae, Echiniscus quadrispinosus, Echiniscus quitensis, Echiniscus rackae, Echiniscus ranzii, Echiniscus reticulatus, Echiniscus reymondi, Echiniscus robertsi, Echiniscus rodnae, Echiniscus rufoviridis, Echiniscus rugospinosus, Echiniscus scabrospinosus, Echiniscus semifoveolatus, Echiniscus shaanxiensis, Echiniscus siegristi, Echiniscus simba, Echiniscus speciosus, Echiniscus spiculifer, Echiniscus spiniger, Echiniscus spinulosus, Echiniscus storkani, Echiniscus succineus, Echiniscus sylvanus, Echiniscus taibaiensis, Echiniscus tamus, Echiniscus tardus, Echiniscus tenuis, Echiniscus tessellatus, Echiniscus testudo, Echiniscus trisetosus, Echiniscus trojanus, Echiniscus tropicalis, Echiniscus tympanista, Echiniscus velaminis, Echiniscus vinculus, Echiniscus virginicus, Echiniscus viridianus, Echiniscus viridis, Echiniscus viridissimus, Echiniscus walteri, Echiniscus weisseri, Echiniscus wendti, Echiniscus zetotrymus.

Hypechiniscus: Hypechiniscus exarmatus, Hypechiniscus gladiator, Hypechiniscus geminus, Hypechiniscus cataractus, Hypechiniscus daedalus, Hypechiniscus flavus.

Mopsechiniscus: Mopsechiniscus franciscae, Mopsechiniscus frenoti, Mopsechiniscus granulosus, Mopsechiniscus imberbis, Mopsechiniscus schusteri, Mopsechiniscus tasmanicus.

Novechiniscus: Novechiniscus armadilloides.

Parechiniscus: Parechiniscus chitonides, Parechiniscus unispinosus.

Proechiniscus: Proechiniscus hanneae.

Pseudechiniscus: Pseudechiniscus alberti, Pseudechiniscus asper, Pseudechiniscus bartkei, Pseudechiniscus beasleyi, Pseudechiniscus bidenticulatus, Pseudechiniscus bispinosus, Pseudechiniscus brevimontanus, Pseudechiniscus clavatus, Pseudechiniscus conifer, Pseudechiniscus dicrani, Pseudechiniscus distinctus, Pseudechiniscus facettalis, Pseudechiniscus goedeni, Pseudechiniscus gullii, Pseudechiniscus insolitus, Pseudechiniscus islandicus, Pseudechiniscus jiroveci, Pseudechiniscus juanitae, Pseudechiniscus jubatus, Pseudechiniscus megacephalus, Pseudechiniscus nataliae, Pseudechiniscus novaezeelandiae, Pseudechiniscus occultus, Pseudechiniscus papillosus, Pseudechiniscus pilatoi, Pseudechiniscus pseudoconifer, Pseudechiniscus pulcher, Pseudechiniscus quadrilobatus, Pseudechiniscus ramazzottii, Pseudechiniscus raneyi, Pseudechiniscus santomensis, Pseudechiniscus scorteccii, Pseudechiniscus shilinensis, Pseudechiniscus sinensis, Pseudechiniscus spinerectus, Pseudechiniscus suillus, Pseudechiniscus transsylvanicus, Pseudechiniscus victor, Pseudechiniscus yunnanensis, Pseudechiniscus mascarenensis.

Testechiniscus: Testechiniscus laterculus, Testechiniscus macronyx, Testechiniscus meridionalis, Testechiniscus spitsbergensis.

Family: Echiniscoididae

Anisonyches: Anisonyches deliquus, Anisonyches diakidius, Anisonyches eleutherensis, Anisonyches mauritianus.

Echiniscoides: Echiniscoides andamanensis, Echiniscoides basalticus, Echiniscoides bruni, Echiniscoides bufocephalus, Echiniscoides costaricensis, Echiniscoides galliensis, Echiniscoides groenlandicus, Echiniscoides hispaniensis, Echiniscoides higginsi, Echiniscoides hoepneri, Echiniscoides horningi, Echiniscoides lichenophilus, Echiniscoides mediterranicus, Echiniscoides musa, Echiniscoides pollocki, Echiniscoides polynesiensis, Echiniscoides porphyrae, Echiniscoides ritavargasae, Echiniscoides rugostellatus, Echiniscoides sigismundi, Echiniscoides travei, Echiniscoides trichosus, Echiniscoides verrucariae, Echiniscoides wyethi.

Family: Oreellidae

Oreella: Oreella chugachii, Oreella mollis, Oreella vilucensis.

(Archechiniscus biscaynei)

It is known only from Biscayne Bay.

(Batillipes adriaticus)

It is known only from the Mediterranean Basin.

(Batillipes mirus)

This is one of the most frequently reported marine species and has been recorded predominantly in the Northern Hemisphere.

(Batillipes lusitanus)

(Batillipes tubernatis)

Body length: ≈219 µm.

It is frequently reported from the waters around the UK, but has also been found off the coast of Florida and Brazil.

(Batillipes noerrevangi)

It is known only from intertidal localities with low water salinity (4–20%) in the North Sea.

(Batillipes pennaki)

This is a common and widely distributed species, but mainly in the temperate zone of the northern hemisphere.

(Batillipes bullacaudatus)

Body length: <126 µm.

It is known mostly from the North American coast of the Atlantic Ocean. The two records from the Atlantic coast of the UK and one uncertain record from the North Sea.

(Batillipes algharbensis)

(Dipodarctus anaholiensis)

It was first described from intertidal beach sand at depths of 30–60 cm in the Hawaiian Islands, but has since been reported primarily as a subtidal species from isolated and disjunct localities in the Northern Hemisphere.

(Moebjergarctus clarionclippertonensis)

The type locality contained in Clarion Clipperton Zone.

(Florarctus heimi)

It was described from the Coral Sea in the shallow subtidal zone, but more recently from the Indian Ocean from quite deep in the subtidal zone.

(Florarctus hulingsi)

More than 80% of records are concentrated in the Mediterranean basin. Outside this area, the species has been reported from tropical localities in the Atlantic and Indian Oceans, as well as from the temperate Celtic Sea.