Tholoarctus

Jørgensen, A., Boesgaard, T.M., Møbjerg, N. & Kristensen, R.M. (2014) The tardigrade fauna of Australian marine caves: With descriptions of nine new species of Arthrotardigrada. Zootaxa, 3802(4): 401–443.

Family: Styraconyxidae

Click taxa for descriptions

Genus description from Kristensen & Renaud-Mornant 1983: “Epicuticule externe en forme de cloche, corps allongé, mince; tête avec assortiment complet d’appendices céphaliques. Clavas primaries ovoïdes, clavas secondaires élargies ventralement. Soies coxales seulement présentes sur P1; P4 avec papille. Cirres A et E avec cirrophore en entonnoir, dans lequel une partie du scapus peut se rétracter. Griffes à trois crochets, les crochets accessoires très fins; mâles et juvéniles pouvant avoir des griffes simples sur les doigts externes. Tube buccal muni de deux excroissances latérales à mi-longueur. Supports de stylets longs et fins et stylets sans articulation sur le tube buccal. Placoïdes réduits ou manquants ; seuls, deux petits apodèmes ronds se trouvent à la partie antérieure du bulbe pharyngien.”

Genus description from Jørgensen et al. 2014: “Styraconyxinae with a barrel-shaped outer epicuticle surrounding the trunk, the body itself is thin and elongated. Complete set of cephalic appendages. Cone-shaped primary clavae and secondary clavae enlarged surrounding both the internal and external cirri. Cirri A and E partially surround by funnel-shaped epicuticle folds. Leg sense organs always present on the first (se1) and fourth (se4) legs, though much smaller sense organs may be present on the second (se2) and third (se3) legs. Flagellum on the sense organ of the first leg always very long. Sense organ of the fourth leg partially enclosed in funnel-shaped epicuticular folds. Females have claws with a main hook, a basal accessory hook and a terminal accessory spur. Males and larvae might have simple claws in the external digits. Long buccal tube with two lateral projections at ca. the mid-point. Long and thin stylet supports articulate with the furcae. Placoids consist of a non-calcified cuticular lining, but three small calcified apodemes occur in the anterior part of the pharyngeal bulb. Mouth cone is terminal and protruding. Globular coelomocytes seem to be attached to the midgut. Sexual dimorphism in body size and claws.”

Fontoura, P., Bartels, P.J., Jørgensen, A., Kristensen, R.M. & Hansen, J.G. (2017) A dichotomous key to the genera of the marine Heterotardigrades (Tardigrada). Zootaxa, 4294: 1–45.
Fontoura, P., Bartels, P.J., Jørgensen, A., Kristensen, R.M. & Hansen, J.G. (2017) A dichotomous key to the genera of the marine Heterotardigrades (Tardigrada). Zootaxa, 4294: 1–45.

Citations:

Jørgensen, A., Boesgaard, T.M., Møbjerg, N. & Kristensen, R.M. (2014) The tardigrade fauna of Australian marine caves: With descriptions of nine new species of Arthrotardigrada. Zootaxa, 3802(4): 401–443.

Kristensen, R.M. & Renaud-Mornant, J. (1983) Existence d’arthrotardigrades semi-benthiques de genres nouveaux de la sous-famille des Styraconyxinae subfam. nov. Cahiers de Biologie Marine, 24: 337–353.

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?

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, focus on other characters first!


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?

Modified from Lindahl K, Balser S.  1999. Key to tardigrade genera [Internet]. Available from: https://sun.iwu.edu/~tardisdp/Keypage33.html