Tardigrades of North Carolina

sciName	taxonomic_comments	body_length	id_comments	habitat	observation_methods	state_protection
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Batillipes pennaki checklist_number: 2245.0	This species appears to be cosmopolitan, but distinct morphological traits were found for Western Atlantic, Eastern Atlantic and Mediterranean populations by Santos et al. 2019 suggesting a pseudocryptic complex. DNA studies have not been conducted on these populations.	av 150-164	Thickset body; eyes not visible after slide mounting; dorsal cuticle finely punctated with poorly marked transverse folds. A small lateral body projection (projection 1) is present between the head an...	Most commonly reported from intertidal beach sand but also collected from shallow subtidal sediments....	DIC or Phase Microscopy....
Batillipes cf. tubernatis checklist_number: 2248.0	Batillipes tubernatis was originally described from Scotland (Pollock 1971), then redescribed based on species from the Gulf of Mexico in Florida by McKirdy (1975). The European populations have no caudal apparatus while 80% of the Florida population did have a caudal apparatus. Santos et al. (2019) concluded that species assigned to B. tubernatis from the Gulf of Mexico and other places outside of Europe are a different species. Thus, we have used the name B. cf. tubernatis to differentiate this species from the European B. tubernatis.	ave. 219 microns (Florid	Either without caudal appendage, or with a strong, single, swollen-based cauidal spike inserting as a continuous extension of the body, or with a single, small off-center caudal spur; lateral body pro...	Intertidal beach sand....	DIC or Phase microscopy....
Batillipes caudatus checklist_number: 2235.0	This is a case of taxonomic intrigue. In 1917, W.P. Hay, a high school biology teacher from Washington, D.C. described the new species Batillipes caudatus from Dictyota on a jetty at Shackleford Bank. Subsequently, Marcus (1929) synonymized B. caudatus with B. mirus. My colleague Paulo Fontoura and I have examined new specimens from North Carolina and original type material, and we believe that Batillipes caudatus and B. mirus are distinct species. Thus, although the current Checklist (Degma & Guidetti 2023) does not officially recognize B. caudatus we include it here to open the case for future investigation. Santos et al. (2018) included B. caudatus in their key to the genus.		Body stout, plump and covered with a finely-granulate almost transparent skin. Head broad, flattened, and bearing seven cirri, one of which is situated on the median line of the top of the head some d...	Dictyota on jetty and intertidal beach sand....	PC, DIC microscopy...
Claxtonia maucci checklist_number: 1990.0	Transferred from Echiniscus by Gasiorek et al. 2019. DNA analysis by Gasiorek et al. 2019 confirmed species identity and included specimens from Roan Mountain, TN.	250 microns maxiumum.	Reddish color; the sculpture is a very coarse and irregular granulation, finer in the anterior part of the paired plates, where it is also partially lacking: it is also absent on a good part of the ce...	Tree lichen and tree moss. Rarely in rock lichen and rock moss....	PC, DIC, and Fluorescence Microsocpy....
Barbaria danieli checklist_number: 1985.0	Transferred from Echiniscus by Gasiorek et al. 2022.	145-204 microns	An Echiniscus of the bigranulatus group, dorsal plates with pores and true granulation, no appendages other than head appendages and lateral cirri A, weak development of dentate collars on the fourth ...	Found on lichen and mosses on trees in urban settings....	PC, DIC and epifluorescence microscopy....
sciName	taxonomic_comments	body_length	id_comments	habitat	observation_methods	state_protection
Echiniscus virginicus checklist_number: 2030.0	Considered a pseudocryptic species with E. lineatus that is pantropical. Only morphological distinction is different cuticle pore structure observable with SEM. TN specimens (and from other eastern US states) positively identified as E. virginicus by DNA analysis (Gasiorek et al. 2019).	up to 183 microns	Length up to 183 microns, sculpture composed of a dense granulation. Presence or absence of eye spots and coloration of the tardigrade not indicated by the author. The internal and external buccal cir...	Tree lichen and tree moss, less commonly in other terrestrial habitats....	DIC and PC microscopy....
Mesobiotus n. species checklist_number: 1830.0	This is a probable new species discovered in the GSMNP. Needs fresh material for complete description.	Unknown.	The genus Mesobiotus is characterized by three roundish macroplacoids in a curved line with a microplacoid very near the end of the 2nd macroplacoid. The species we find in the Smokies and surrounds h...	Predominantly soil/leaf litter. Also found less commonly in all other habitat types....	DIC and PC....
Murrayon hastatus checklist_number: 1950.0	Morphologically indistinguishable from species originally described from Europe, no molecular evidence for further comparison. Eggs needed for positive ID.	Up to 312 microns.	This is an aquatic species, very wide-spread for example in moss of peat-bogs. Length up to 312 µm, eye spots present. Buccal tube medium wide (diameter about 4.5 µm), pharynx oval contain...	Freshwater streams....	PC and DIC....
checklist_number: 2025.0	This species was redescribed and nucleotide sequences were added to Genbank by Kaczmarek et al. 2022 from populations in Germany. The particular subspecies reported from NC is no longer considered valid. Molecular data is not available for N.Am. specimens.	Up to 280 microns.	Females (measurements and statistics in Table 1, Supp. fi le 2 and Figs 1–4). Body orange in live specimens and transparent/light yellow after preparation. Eyes red. Apart from the head appendag...	Moss...	DIC, PC, and SEM....
Hypsibius n. species checklist_number: 1280.0	Probable new species from GSMNP.	Unknown.	Need fresh material for complete description....	Stream periphyton and sediments....	DIC and PC....
sciName	taxonomic_comments	body_length	id_comments	habitat	observation_methods	state_protection
Dianea basalovoi checklist_number: 1670.0	Transfered from Isohypsibius to Dianea by Gasiorek et al 2019. Morphologically indistinguishable from species reported from Europe but molecular evidence for further comparison unavailable.	To 280 microns.	Colorless. Eye spots present. The sculpture of the cuticle is composed of irregular polygonal plates, of variable size, which form a very pronounced network (maximum size of the mesh about 5 µm)...	Freshwater streams and periphyton. Rarely see in terrestrial samples....	PC, DIC...
Batillipes bullacaudatus checklist_number: 2230.0	The genus Batillipes is the most speciose of all marine tardigrade genera, containing almost 40 species (about 16% of all marine tardigrade species).	up to 126 microns	Median cirus (20 microns) lacking base, internal buccal cirri 20 microns) with base; external buccal cirri (14 microns), inserted on the cephalic papillae; clavae (13 microns); lateral cirri (28~) ori...	Intertidal sand. In collections on Pawley's Island, SC, much more common on sound-side than ocean-side....	PC, DIC, and fluorescent microscopy....
Milnesium n. species checklist_number: 1030.0	Probable new species from the Warren Wilson College campus appears to be the same species found in the Smokies. This species is now under active investigation by Bartels & Kaczmarek.	436-696 microns.	Description in progress....	Tree moss and lichen....	DIC, PC, SEM, and DNA analysis....
Viridiscus perviridis checklist_number: 2130.0	The genus was recently analyzed with integrative taxonomy by Momeni et al. 2023. This species was transferred from Echiniscus by Gasiorek et al. 2019.	185-285 microns.	In reflected light the tardigrade appears dorsally as a very dark brownish-green color, almost black, while ventrally the color is yellowish in the young and brick-red in the adults; in transmitted li...	Moss....	PC and DIC....
Hypechiniscus gladiator group checklist_number: 2060.0	This record is probably Hypechiniscus daedalus. At the time of this study H. gladiator was considered to be cosmopolitan. Gasiorek et al (2021) used integrative taxonomy to untangle a pseudocryptic cluster of species and described the new species H. daedalus and concluded that it is likely endemic to N. America. Hypechiniscus gladiator is verified only in Scotland and Iceland, thus we have changed this record to H. gladiator group.	166-300 microns	For H. gladiator proper: Females (i.e. from the third instar onwards; measurements and statistics in Table 4): Body whitish and stout (Figs 3A, C, 4), with spheroid or ovoid black eyes, persisting al...	Moss and lichen and other terrestrial habitats....	PC/DIC/SEM...
sciName	taxonomic_comments	body_length	id_comments	habitat	observation_methods	state_protection
Viridiscus viridianus checklist_number: 2140.0	The genus was recently analyzed with integrative taxonomy by Momeni et al. 2023. Species transferred from Echiniscus by Gasiorek et al. 2019. Species description amended by Momeni et al. 2023.	220-336 microns.	Color green (U.S.A.) or dark grey (Azores); eye spots present; plate margins well marked, median plate 1 triangular and distant from scapular plate; median plate 3 absent but the area between paired p...	Rock lichen. Also found in one bird nest....	PC and DIC....
Viridiscus celatus checklist_number: 2125.0	The genus was recently analyzed with integrative taxonomy, and this new species was described from Johnson City, TN by Momeni et al. 2023. A reanalysis of specimens from GSMNP previously described as Viridiscus perviridis led to new species designation.	152-252 microns.	Females (i.e., from the third instar onwards; measurements and statistics in Table 6). Body medium sized and bulky. Body cavity with yellowish pigments (typical for most echiniscids), whereas dorsal a...	Moss....	PC and DIC....
Viridiscus unidentified species checklist_number: 2170.0	Transferred from Echiniscus by Gasiorek et al. 2019. Species description for V. viridis amended by Pilato et al. 2008.	190-220 microns.	For the actual Viridiscus viridis: The second specimen about 220 μm long. Colour of plates dark green. No true neck plate, but a thickened transversal area with a faint ornamentation is present be...	Moss and lichen....	PC....
Viridiscus viridissimus checklist_number: 2160.0	The genus was recently analyzed with integrative taxonomy by Momeni et al. 2023. This species was transferred from Echiniscus by Gasiorek et al. 2019. A newly described species Viridiscus miraviridis Nelson, Adkins Fletcher, Guidetti, Roszkowska, Grobys & Kaczmarek, 2021 was synonymized with V. viridissimus by Gasiorek et al. 2021.	180 microns.	Color olive green, not only of the plates, but of all the cuticle of the body, including the ventral surface, the legs, the claws, and the appendices: the plates, the claws, and the dentate collars of...	In GSMNP rock lichen and one bird nest. In Johnson City, TN, Nelson found it on moss on concrete (limestone) exposed to sun and wind....	DIC and PC....
Pilatobius n. species checklist_number: 1380.0	This is a presumptive new species discovered in the GSMNP. It is very similar to P. burti, but cuticle with punctation.	Unknown.	Needs further study....	Lichen on Frazier fir tree....	PC and DIC....
sciName	taxonomic_comments	body_length	id_comments	habitat	observation_methods	state_protection
Ursulinius lunulatus checklist_number: 1740.0	Transferred from Isohypsibius by Gasiorek et al. 2019, nomen inquirendum according to Gasiorek et al. 2019 due to insufficient original description. Morphologically indistinguishable from species first described in Hungary, no DNA for further comparison.	150-280 microns.	Length up to 280 µm, colorless, eye spots present. Dorsally, on its flanks and on the basal portion of the legs, the cuticle is sculptured with big rounded granules or, sometimes, polygons (quad...	Soil/leaf litter. Rarely in moss and stream samples....	PC and DIC....
Thulinius augusti checklist_number: 1630.0	Transferred to Thulinius and species redescribed by Bertolani et al. 1999. The genus was transferred to Doryphoribiidae by Gasiorek et al. 2019.	Unknown.	Cuticle smooth and without pigmentation. Eye spots are absent in the mounted material. The buccal aperture is surrounded by 6 lobes usually subdivided into 12-16 sublobes, of various shapes and size...	Freshwater streams....	PC and DIC....
Tholoarctus natans natans checklist_number: 2320.0	This genus contains only two species. Tholarctus natans is divided into two subspecies, T. natans natans and T. natans pedunculatus.	110-170 microns.	External epicuticle bell-shaped, body elongated, thin: complete complement of cephallic appendages. Primary clavae ovoid, secondary clavae expanded ventrally. Coxal setae (leg sense organs) only pres...	Deep sea fine sand....	Unknown....
Tanarctus gracilis checklist_number: 2200.0	There are currently (Oct 2023) 15 species known for this genus.	80 microns.	Body oval, the head is trapezoidal, with lateral border posteriorly curved; the anterior part of the head is formed from a cuticular membrane, which unites the bases of the internal median cirri. The ...	Deep sea fine sand....	PC....
Stygarctus bradypus checklist_number: 2280.0	There are currently (October 2023) eight species in this genus.	90-150 microns.	Stygarctus bradypus proper: The body appears clearly segmented into 5 parts: a cephalic region (plate) well delimited, three unpaired dorsal plates, which also encompass the body laterally as far as ...	Intertidal beach sand....	PC. ...
sciName	taxonomic_comments	body_length	id_comments	habitat	observation_methods	state_protection
Ramazzottius n. species checklist_number: 1510.0	Probable new species from GSMNP. Need fresh material for full analysis.	Unknown.	Needs further analysis....	Tree lichen. Also soil and tree moss....	PC and DIC....
Pseudechiniscus (Pseudechiniscus) n. species 2 checklist_number: 2100.0	This is a probable new species from the GSMNP. See Pseudechiniscus suillus species account for taxonomic details about this group.	Unknown.	Further work is needed....	Moss and lichens....	PC and DIC....
Ramazzottius cf. oberhaeuseri checklist_number: 1520.0	This is a pseudocryptic species complex that was examined and an amended species description given by Stec et al. 2018.	Maximum 500 microns.	Pigmentation of the cuticle from a light-red to a red-brown chequered pattern on the dorsum (Fig. 3). The pigmented surface is divided by four longitudinal and eight transversal transparent stripes th...	Moss and lichen....	PC....
Platicrista angustata checklist_number: 1430.0	Morphologically indistinguishable from species originally described in Scotland. Records for this species exist in GenBank, but there is no molecular data for the NC population for further comparison. Family designation follows Tumanov & Tsvetkova (2023).	350-490 microns.	Transparent, with smooth cuticle, eyes absent. The body has its maximum width (observed from dorsum) in correspondence with the 3rd pair of legs, then reducing progressively toward the rostral end, be...	Soil/leaf litter, less often in tree moss and lichen....	PC and DIC....
Paramacrobiotus tonollii checklist_number: 1910.0	Transferred from Macrobiotus by Guidetti et al. 2009. Part of the P. areolatus complex with 3 rows of macroplacoids and no microplacoid.	Up to 620 microns.	Length up to 620 µm, colorless (white in reflected light), eyes present. The cuticle is smooth, but presents dorsally numerous small “pearls”, clearly visible only with immersion obj...	Tree lichen and moss. Found less commonly in all other habitat types....	DIC and PC....
sciName	taxonomic_comments	body_length	id_comments	habitat	observation_methods	state_protection
Paramacrobiotus halei checklist_number: 1890.0	New species discovered in GSMNP. Transferred to Paramacrobiotus by Bartels & Nelson 2012. Part of the P. richtersi group, containing 3 macroplacoids and a microplacoid.	283-450 microns.	Colorless, cuticle with very small tubercles, quite difficult to see; very fine granules present on all legs, more evident on legs IV; eyes absent. Buccal armature in oral cavity of richtersi-type; bu...	Soil / leaf litter, less commonly in other terrestrial habitats. ...	DIC and PC. ...
Paramacrobiotus aff. areolatus checklist_number: 1880.0	Transferred from Macrobiotus by Guidetti et al. 2009, amended by Stec et al. 2020. Paramacrobiotus areolatus is part of the P. areolatus species complex with three rows of macroplacoids and no microplacoid. The complex has been separated into several genetically distinct species (Stec 2020).	Up to 800 microns.	For Paramacrobiotus areolatus s.s.: In live animals, body almost transparent in juveniles and white in adults; after fixation in Hoyer’s medium body transparent (Fig. 1A). Eyes present in liv...	Moss....	DIC and PC....
Minibiotus intermedius checklist_number: 1850.0	This species was integratively redescribed and the neotype was designated by Kaczmarek et al. 2022.	149-245 microns.	The body cuticle was smooth without pores and granulation, but very poorly visible granulation on the IV pair of legs was present. Large eye spots were also present. The buccal tube with an anterior a...	Tree moss. Less commonly in soil, and rarely in othe terrestrial habitats....	DIC, PC. ...
Milnesium bohleberi checklist_number: 1010.0	New species discovered in GSMNP. Named in honor of Steve Bohleber, long time friend of the Smokies. DNA data is available for specimens from Gregory's Cave, GSMNP, USA. (Morek et al. 2021).	620-880 microns.	Body white before fixation and transparent afterwards, eyes present. Cuticle smooth, without granulation, pseudopores, pores, reticulations, or gibbosities. Six peribuccal papillae (ventral papilla sm...	Tree moss and lichen....	DIC and PC....
Microhypsibius bertolanii checklist_number: 1460.0	Morphologically indistinguishable from species first described in Greenland. DNA sequences are available in GenBank, but fresh specimens from the smokies are necessary for further comparison.	To 143 microns.	Smooth cuticle, eyes absent. The forehead is strongly bulging, with anterio-ventral buccal aperture. Mouth without lamellae or papillae. Buccal tube rather slender, with a thin reinforcement bar. Phar...	Freshwater streams....	PC and DIC....
sciName	taxonomic_comments	body_length	id_comments	habitat	observation_methods	state_protection
Mesocrista spitzbergensis checklist_number: 1310.0	Morphologically matches species from Svalbard amended by Gasiorek et al. 2016 with integrative taxonomy. No DNA data for smokies population. Family designation follows Tumanov & Tsvetkova (2023).	399-548 microns.	Genus diagnosis: Peribuccal structures absent. Apophyses for the insertion of stylet muscles in the shape of wide and flat ridges symmetrical with respect to the frontal plane and with welldeveloped ...	Soil and tree moss....	DIC and PC....
Macrobiotus nelsonae checklist_number: 1800.0	Currently no molecular data is available for this species.	230-925 microns.	Animals of large size. Smooth cuticle with pores. Eye-spots anterior. Wide buccal tube; evident buccal armature, with two consistent bands of teeth and with transverse crests. Two rod-shaped macro...	Soil / leaf litter. Rarely found in stream sediment....	DIC and PC....
Macrobiotus margoae checklist_number: 1780.0	This is a new species discovered in the GSMNP. The species was delineated from the Macrobiotus pallari complex using integrative taxonomy by Stec et al. 2021. It is named in honor of Margo Nottoli, Paul Bartels' life partner, as a partial thank you for her infinite (?) tolerance of his tardigrade obsession.	239-443 microns.	In live animals, body almost transparent in smaller specimens and whitish in larger animals; transparent after fixation in Hoyer’s medium (Fig. 30). Eyes present in live animals and after fixati...	Tree moss and lichen, and also in all other terrestrial habitats....	PC and DIC....
Macrobiotus hufelandi checklist_number: 1760.0	Macrobiotus hufelandi was once thought to be cosmopolitan, but it is now known to be a very large group of difficult to identify species. Kaczmarek & Michalczyk 2017 provides valuable morphological guidance for this group.	Unknown.	The hufelandi group have two macroplacoids and a microplacoid. In Macrobiotus hufelandi proper the dorsal cuticle is smooth without gibbosities or granulations, oral cavity armature of hufelandi type ...	Most abundant in soil, but also found in all other terrestrial habitats....	DIC and PC....
Macrobiotus hibiscus checklist_number: 1755.0	Nomen inquirendum according to Stec et al. 2021 due to insufficient detail in description for positive identification. Very similar to others in M. hufelandi group.	Up to 340 microns.	Colorless, eyespots present. Mouth opening consisting of 3 rings (the author does not cite peribuccal lamellae); buccal tube rather short (30 microns) and thin (2.5 microns); pharynx almost round, wit...	Moss....	PC and DIC....
sciName	taxonomic_comments	body_length	id_comments	habitat	observation_methods	state_protection
Ligiarctus eastwardi checklist_number: 2260.0	This is the only species in the genus.	approimately 120 microns	A genus of Florarctinae with narrow head. Wide implementation of the primary clavae occupying the lateral edge of the head. Large, rigid and backwards bending primary clavae (shared with Higginsarctus...	Deep sea fine sand, 400 m bsl. ...	DIC and PC....
Hypsibius n. species (aff. dujardini) checklist_number: 1290.0	This is a probable new species from the GSMNP, similar to Hysibius dujardini.	Unknown.	Needs fresh material for a complete description....	Soil....	PC and DIC....
Hypsibius convergens checklist_number: 1260.0	Probably a species complex in need of integrative taxonomic examination.	Up to 400 microns.	Length up to 400 µm; eyes present, sometimes small, sometimes of great size. Buccal tube rather narrow, from 1.5 to 1.7 µm in diameter, with appendices of insertion for the muscles in the ...	Soil / leaf litter, tree moss, tree lichens, rock moss, also occurs in streams....	DIC, PC....
Hypechiniscus daedalus checklist_number: 2050.0	This is a new species from the GSMNP. The species was recently described with integrative taxonomy methodology by Gasiorek et al. 2021. Type locality is Purchase Knob, GSMNP. Old records of H. gladiator from N. Am. are likely to be this new species.	180-240 microns	Females: Body whitish and stout (Figs 3A, C, 4), with spheroid or ovoid black eyes, persisting also after mounting. Elongated, dactyloid cephalic papillae (secondary clavae) and (primary) clavae (Figs...	Tree moss and less often tree lichen. Rarely in rock moss and soil....	DIC, PC, and SEM....
Guidettion prorsirostre checklist_number: 1240.0	Transferred from Adropion by Gasiorek & Michalczyk 2020. Morphologically matches European populations, but no molecular data for further comparison. Family designation follows Tumanov & Tsvetkova (2023).	245-325 microns.	Body slender, buccal aperture terminal as in the genus Macrobiotus, so that the “forehead” becomes receding and all the cephalic region, viewed in profile, assumes an appearance more of Ma...	Terrestrial in European collections, freshwater stream in the one Smokies record....	DIC and PC....
sciName	taxonomic_comments	body_length	id_comments	habitat	observation_methods	state_protection
Bertolanius smreczynskii checklist_number: 1050.0	Morphologically inidstinguishable from species first described from Axel Heiberg Island, Nunatuk, Canada. No molecular evidence for further comparison.	up to 969 microns	Cuticle smooth. In incident light olive-green colour of individuals. Coloured cuticle and body cavity. Black pigment of eyes. Sucking pharynx possessing three barlike macroplacoids, the first of which...	Rock moss and soil/leaf litter....	PC and DIC microscopy....
Grevenius granulifer checklist_number: 1580.0	Transferred from Isohypsibius by Gasiorek et al. 2019. Morphologically identical to European specimens, but no molecular data for further comparisons.	Up to 405 microns.	Length up to 405 µm. The body is white. Cuticle covered with small thickenings (up to 1.5 µm), irregularly distributed (Pl. XVI d), being the biggest in the middle and the rear of the body...	Freshwater streams....	PC and DIC...
Doryphoribius n. species checklist_number: 1560.0	This is a presumptive new species from the GSMNP. It needs fresh material for a full description.	Unknown.	Needs full description....	Freshwater stream periphyton....	PC and DIC....
Doryphoribius longistipes checklist_number: 1550.0	This is a new species discovered in Bartels' Smokies inventory, described in Bartels et al. 2008.	Up to 280 microns.	Body length 298.0 µm. Body colour unknown, eyes present. Cuticle smooth. Mouth anteroventral. Bucco-pharyngeal apparatus of the Doryphoribius type: peribuccal lamellae and papulae absent, ventra...	Freshwater streams in periphyton in NC samples, in TN found in wet limestone cave sediments in Cades Cove....	PC and DIC...
Doryphoribius doryphorus checklist_number: 1540.0	Morphologically indistinguishable from species originally discovered in Sicily. No molecular evidence for further comparisons.	Maximum 200 microns.	Small, with maximum size of 200 µm, colorless, smooth cuticle and eyes in posterior position. Ventral mouth, without peribuccal lamellae; forehead sloped. Buccal tube with reinforcement bar, 25 ...	Stream sediment....	PC and DIC microscopy....