Hebesuncus

Class : Eutardigrada

Order : Hypsibioidea

Family : Ramazzottiidae

Genus : Hebesuncus

Genus Hebesuncus

Hypsibioidea from Pilato 1969 in Marley et al. 2011: “Parachela; claws asymmetrical (2121); Hypsibius-type claw pairs; AISM hooked (or, if the buccal tube is elongated, AISM can be broad ridges).”

Hypsibioidea from Bertolani et al. 2014: “Double claws asymmetrical with respect to the median plane of the leg (2121), the external (or posterior) claw often with flexible main branch; double claws different in size and shape on the same leg (Hypsibius and Ramazzottius type, or modified), or very reduced in size (Calohypsibius and Microhypsibius type); buccal tube often very narrow”

Hypsibioidea from Gąsiorek et al. 2018: “Eutardigrades with asymmetrical claws (2-1-2-1) and pseudolunulae at claw bases or without any cuticular structures under the basal parts. Hooked or broad-ridged apophyses for the insertion of the stylet muscles. Herbivorous or microbivorous (Guidetti et al. 2012).”

Ramazzottiidae from Marley et al. 2011: “Hypsibioidea. The AISM comprises asymmetric, dissimilar dorsal and ventral ‘blunt hooks’. Claw pairs asymmetric (2121), external claw primary branch joins the secondary claw and basal section with flexible junction; primary branch may be very long and slender; the internal claw branches and basal section unified into a single rigid element. Eggs have a sculptured chorion and are deposited free in the environment.”

Ramazzottiidae from Bertolani et al. 2014: “Double claws different in shape and size on the same leg, the external (or posterior) claw with the primary branch connected to basal tract with an evident, thin, flexible tract (Ramazzottius or oberhaeuseri type, or modified), or completely disconnected. Eggs laid freely and with ornamented shell.”

Genus description from Pilato 1987: “The bucco-pharyngeal apparatus is of the Diphascon model with the apophyses for the insertion of the muscles of the stylets in the shape of a ‘blunt hook’; the dorsal hook is different in shape and size from the ventral one with consequent asymmetry with respect to the frontal plane; the caudal processes of the apophyses for the insertion of the muscles of the stylets are pointing backwards and sideways; the furcae have the postero-lateral processes thickened at their apices.

Other characters: The bucco-pharyngeal tube is narrow; the pharyngeal tube is very short, almost half as long as the pharyngeal bulb and clearly shorter than the buccal tube; the ‘drop-like’ thickening is absent; the stylet supports are inserted just after 2/3 of the buccal tube length; the pharyngeal apophyses are well developed; the pharyngeal bulb is rounded with short placoids.”


Citations :

Hebesuncus image from Dastych H, JM. 1994. A new species of the genus Hebesuncus from the Antarctic nunataks, Dronning Maud Land (Tardigrada). Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg. 11 (150): 139-145.

“Parts” images from Pilato G, Binda MG. 2010. Definition of families, subfamilies, genera, and subgenera of the Eutardigrada, and keys to their identification. Zootaxa. 2404: 1-54.

Bertolani R, Guidetti R, Marchioro T, Altiero T, Rebecchi L, Cesari M. 2014. Phyloeny of Eutardigrada: New molecular data and their morphological support lead to the identification of new evolutionary lineages. Molecular Phylogenetics and Evolution. 76: 110-126.

Gąsiorek P, Stec D, Morek W, Michalczyk Ł. 2018. An integrative redescription of Hypsibius dujardini (Doyère, 1840), the nominal taxon for Hypsibioidea (Tardigrada: Eutardigrada). Zootaxa. 4415 (1): 45-75.

Marley NJ, McInnes SJ, Sands CJ. 2011. Phylum Tardigrada: A re-evaluation of the Parachela. Zootaxa. 2819: 51-64.

Pilato G. 1969. Schema per una nuova sistemazione delle famiglie e dei generi degli Eutardigrada. Bollettino delle Sedute della Accademia Gioenia di Scienze Naturali in Catania, Series IV. 10 (277): 181-193.

Pilato G. 1987. Revision of the genus Diphascon Plate, 1889, with remarks on the subfamily Itaquasconinae (Eutardigrada, Hypsibiidae). pp. 337-357 in Bertolani R (ed). Biology of Tardigrades: Selected symposia and monographs.

Claw Symmetry Relative To Median Plane Of Leg?

Image from Bingemer J, Hohberg K. 2017. An illustrated identification key to the eutardigrade species (Tardigrada, Eutardigrada) presently known from European soils. Soil Organisms. 89 (3): 127-149.

Stylet Support Insertion Point As Percentage Of Buccal Tube Length

Stylet support insertion point = ss divided by Buccal tube length, as %. Note anterior measurements begin at anterior margin of stylet sheaths, ss is centred where stylet supports reach buccal tube (this requires a good dorsal or lateral view for proper measurement)


Image from Tumanov DV. 2006. Five new species of the genus Milnesium (Tardigrada, Eutardigrada, Milnesiidae). Zootaxa. 1122: 1-23.

Buccal Tube Long, Mouth At Anterior Of A Protrusible Snout

Images from Pilato G, Binda MG. 2010. Definition of families, subfamilies, genera, and subgenera of the Eutardigrada, and keys to their identification. Zootaxa. 2404: 1-54.

Pharyngeal Tube With Spiral Reinforcement

Images from Pilato G, Binda MG. 2010. Definition of families, subfamilies, genera, and subgenera of the Eutardigrada, and keys to their identification. Zootaxa. 2404: 1-54.

Peribuccal papillae?

Note: in a tardigrade with snout extended, the flex of the pharyngeal tube may not be obvious.


Images from Pilato G, Binda MG. 2010. Definition of families, subfamilies, genera, and subgenera of the Eutardigrada, and keys to their identification. Zootaxa. 2404: 1-54.

(Lateral) cephalic papillae, peribuccal papillae, Milnesiidae - type claws, exceptionally wide (and usually short) buccal tube?

Cephalic image modified from Nelson DR, Guidetti R, Rebecchi L. 2009. Tardigrada. Ch. 14 in Ecology and Classification of North American Freshwater Invertebrates, Thorp JH, Covich AP (eds.), 3rd ed.

Claw image modified from Pilato G, Binda MG. 2010. Definition of families, subfamilies, genera, and subgenera of the Eutardigrada, and keys to their identification. Zootaxa. 2404: 1-54.

Pharyngeal tube flexible?

Note: in a tardigrade with snout extended, the flex of the pharyngeal tube may not be obvious.


Images from Pilato G, Binda MG. 2010. Definition of families, subfamilies, genera, and subgenera of the Eutardigrada, and keys to their identification. Zootaxa. 2404: 1-54.

Serrated / dentate cuff / collar on legs IV

Left image from Kristensen RM. 1987. Generic revision of the Echiniscidae (Heterotardigrada), with a discussion of the origin of the family. pp. 261-335 in Bertolani R (ed). Biology of Tardigrades: Selected symposia and monographs.

Right Image from Richters F. 1926. Tardigrada. in Krumbach T. 1927. Handbuch der Zoologie, 3rd band, Walter de Gruyter & Co.

Leg sensory structures

Typically papilla on leg IV (se4 in image) and variable shape on leg I


Image from Kristensen RM. 1987. Generic revision of the Echiniscidae (Heterotardigrada), with a discussion of the origin of the family. pp. 261-335 in Bertolani R (ed). Biology of Tardigrades: Selected symposia and monographs.

Trunk appendages

Appendages (after Cirrus A) labeled according to plate … B (Scapular plate), C (First segmental plate), D (Second segmental plate), E (terminal plate). “B” is lateral, “Bd” is dorsal

Be careful! There can be great variability within a population, with individuals (especially juveniles) lacking some appendages. When in doubt, choose “Unclear” to ignore this character!


Left image from Ramazzotti G, Maucci W. 1983. Il phylum Tardigrada(III edizione riveduta e aggiornata). English translation by C. W. Beasley, 1995. Memorie dell’ Istituto Italiano di Idrobiologia 41: 1-1012.

Right image cropped from Kristensen RM. 1987. Generic revision of the Echiniscidae (Heterotardigrada), with a discussion of the origin of the family. pp. 261-335 in Bertolani R (ed). Biology of Tardigrades: Selected symposia and monographs.

Cirrus A form

Images modified from Kristensen RM. 1987. Generic revision of the Echiniscidae (Heterotardigrada), with a discussion of the origin of the family. pp. 261-335 in Bertolani R (ed). Biology of Tardigrades: Selected symposia and monographs.

Lower image (Mopsechiniscus) modified from du Bois-Reymond Marcus E. 1944. Sobre tardigrados brasileiros. Communicaciones Zoologicas del Museo de Historia Natural de Montevideo. 1(13): 1-19 plus plates.

Terminal / caudal plate with two notches?

Images modified from Kristensen RM. 1987. Generic revision of the Echiniscidae (Heterotardigrada), with a discussion of the origin of the family. pp. 261-335 in Bertolani R (ed). Biology of Tardigrades: Selected symposia and monographs.

First image (Echiniscus) modified from Richters F. 1926. Tardigrada. in Krumbach T. 1927. Handbuch der Zoologie, 3rd band, Walter de Gruyter & Co.

Inner & outer buccal cirri present

Image modified from Nelson DR, Guidetti R, Rebecchi L. 2009. Tardigrada. Ch. 14 in Ecology and Classification of North American Freshwater Invertebrates, Thorp JH, Covich AP (eds.), 3rd ed.

Pseudosegmental plate between last medial plate & terminal plate?

Note on 2nd image, what you see before the terminal plate is either no plate, or a full-width medial plate; don’t confuse the latter with a pseudosegmental plate!
Also, sometimes the pseudosegmental plate is paired, like segmental plates II & III. It’s still pseudosegmental!

Images from Clifford HF. 1991. Aquatic invertebrates of Alberta. University of Alberta Press, Alberta, Canada.

Segmental plates larger than intersegmental?

Image modified from Lindahl & Balser (1999), http://www.iwu.edu/~tardisdp/Keypage35.html

Any Paired Plates?

Images from Kristensen RM. 1987. Generic revision of the Echiniscidae (Heterotardigrada), with a discussion of the origin of the family. pp. 261-335