Doryphoribius

Class : Eutardigrada

Order : Isohypsibioidea

Family : Doryphoribiidea

Genus : Doryphoribius

Genus Doryphoribius

Isohypsibioidea from Marley et al. 2011: “Parachela. Claws asymmetrical (2121); Isohypsibius-type claw pairs; AISM ridged.”

Isohypsibioidea from Bertolani et al. 2014: “Double claws asymmetrical with respect to the median plane of the leg (2121), normally with similar shape and size on each leg; double claws of the Isohypsibius type (secondary branch of the external claw inserted perpendicularly on the claw basal tract), or reduced from it: Hexapodibius type (very short, without common basal tract, with a base as large as the sum of the primary and secondary branch widths, and with an evident suture between primary and secondary branch); Haplomacrobiotus type (one branch only); completely absent (Apodibius). Buccal tube completely rigid (apart Paradiphascon; see below) and often relatively large, without (Dastychius, Eremobiotus, Halobiotus, Isohypsibius, Paradiphascon, Pseudobiotus, Thulinius) or with (Apodibius, Doryphoribius, Haplomacrobiotus, Haplohexapodibius, Hexapodibius, Parhexapodibius) ventral lamina. Eggs with smooth shell laid within the exuvium.”

Isohypsibiidae from Marley et al. 2011: “Isohypsibioidea. Claw pairs of similar size and shape. External and internal claws exhibiting articulation (the basal section and secondary branch form a solid unit while the primary branch and secondary branch articulate). Claws Isohypsibius-type, forming a right-angle between basal section and secondary branch. AISM ridge-like.”

Genus description from Pilato 1969: NOT AVAILABLE, NEED TO GET THIS PAPER

Genus description from Schuster et al. 1980: “Without peribuccal papulae or buccal lamellae; siz peribuccal lobes prsent. Buccal tube short, rigid, without spiraled thickenings; mucrones present. Pharynx with apophyses and macroplacoids; without microplacoids or septulum. Cuticle without pores. Two double claws of each leg of similar size and shape; not connected; without lunulae; sequence 2,1,2,1. Smoooth eggs deposited in cuticle.”

Genus description from Ramazzotti & Maucci: “Hypsibiidae with Isohypsibius type claws; there is present the reinforcement bar of the buccal tube.”

Groups, from Michalczyk & Kaczmarek, 2010
evelinae: dawkinsi, dupliglobulatus, evelinae, flavus, huangguoshuensis, maranguensis, picoensis, quadrituberculatus, zyxiglobus
doryphorus: doryphorus, koreanus, macrodon, neglectus, pilatoi, polynettae, smokiensis, tergumrudis
vietnamensis: gibber, mariae, minimus, vietnamensis
zappalai: bertolanii, korganovae, longistipes, mexicanus, qinlingense, taiwanus, turkmenicus, zappalai

Key from Michalczyk & Kaczmarek, 2010: missing n sp 5

1. Two macroplacoids in the pharynx …………………………………………………………………………………………………………… 2
-. Three macroplacoids in the pharynx………………………………………………………………………………………………………… 3
2(1). Cuticular gibbosities present………………………………………………………………………………………….. 4 (evelinae group)
-. Cuticular gibbosities absent ………………………………………………………………………………………12 (doryphorus group)
3(1). Cuticular gibbosities present…………………………………………………………………………………… 19 (vietnamensis group)
-. Cuticular gibbosities absent ………………………………………………………………………………………….. 22 (zappalai group)
The evelinae group:
4(2). Cuticular gibbosities present only on legs …………………………………………………………………………………………………. 5
-. Cuticular gibbosities present on dorsal side of the body …………………………………………………………………………….. 6
5(4). Dorsal cuticle smooth ……………………………………………………………………………………………………………….D. evelinae
-. Dorsal cuticle with reticular design ………………………………………………………………………………………………..D. flavus
6(4). Cuticular gibbosities indistinct and irregularly arranged …………………………………………………………………………….. 7
-. Cuticular gibbosities distinct and arranged in defined transverse rows ………………………………………………………… 8
7(6). Cuticular tubercles present, lunules absent, hind claws larger than claws I–III ……………………….. D. maranguensis
-. Cuticular tubercles absent, lunules present, hind claws slightly smaller than claws I–III ………………… D. picoensis
8(6). Four dorsal gibbosities arranged in two caudal rows (i.e., configuration: II:2-2) ………………D. quadrituberculatus
-. More than four dorsal gibbosities present, gibbosities arranged in more than two rows and located throughout the dorsal cuticle ……………………………………………………………………………………………………………………………………….. 9
9(8). Seven rows of gibbosities with only two gibbosities in each row (i.e., configuration: VII:2-2-2-2-2-2-2) ………….. ………………………………………………………………………………………………………………………………….. D. dupliglobulatus
-. More than seven rows of gibbosities and with up to six gibbosities per row………………………………………………… 10
10(9). A single gibbosity on each of the hind legs, six gibbosities in row I (gibbosity configuration: IX:6-6-4-6-4-6-4-4- 2+2[LIV]) …………………………………………………………………………………………………………………D. dawkinsi sp. nov.
-. No gibbosities on hind legs, four gibbosities in row I ……………………………………………………………………………….. 11
11(10). Gibbosity configuration IX:4-4-4-4-4-4-4-2-2 ……………………………………………………………… D. huangguoshensis
-. Gibbosity configuration IX:4-6-4-6-4-6-4-4-2 ………………………………………………………………………… D. zyxiglobus
The doryphorus group:
12(2). Claws IV reduced and clearly smaller than claws I–III …………………………………………………………………. D. pilatoi
-. Claws IV similar in size or larger than claws I–III…………………………………………………………………………………… 13
13(12). Entire cuticle without sculpture……………………………………………………………………………………………………………. 14
-. Cuticle at least partially sculptured………………………………………………………………………………………………………… 17
14(13). Lunules on internal claws I–III present, on external absent ………………………………………………………… D. koreanus
-. Internal and external claws without lunules ……………………………………………………………………………………………. 15
15(14). Stylet furcae robust, triangular in shape, with the postero-lateral processes rounded but not swollen D. neglectus
-. Stylet furcae with swollen postero-lateral processes …………………………………………………………………………………. 16
16(15). Oral cavity armature composed of small, numerous teeth in the second band, and one large dorso-median tooth and two less ventro-median teeth in the third band, buccal tube 3.0 µm or more in width …………….. D. macrodon
-. Oral cavity armature absent/not visible under LM, buccal less than 3.0 µm in width …………………..D. doryphorus
17(13). Dorsal cuticle smooth but large granules present on legs IV …………………………………………………. D. tergumrudis
-. Dorsal cuticle sculptured, granules on legs IV absent ……………………………………………………………………………….. 18
18(17). Ventral sculpture absent, lunules on claws absent, pt of buccal tube external width more than 20, eyes absent ….. ………………………………………………………………………………………………………………………………………… D. smokiensis
-. Ventral sculpture present, lunules on claws present, pt of buccal tube width less than 15, eyes present ……………… ………………………………………………………………………………………………………………………………………… D. polynettae
The vietnamensis group:
19(3). Eight rows of gibbosities (configuration: VIII:2-2-2-2-2-2-2-2), eyes absent ………………………………… D. minimus
-. More than eight rows of gibbosities, eyes present…………………………………………………………………………………….. 20
20(19). Nine rows of gibbosities (configuration: IX:5-4-5-4-5-4-5-2-3), dorsal cuticle with fine granulation ……………….. ……………………………………………………………………………………………………………………………………… D. vietnamensis
-. More than nine rows of gibbosities, different gibbosity configuration, cuticle with polygonal tubercles ………… 21
21(20). Ten rows of well visible gibbosities (configuration: X:4-4-4-4-4-4-4-2-2-3), undulations absent, ……….D. gibber
-. Twelve rows of poorly visible gibbosities (configuration: XII:2-4-2-4-2-4-2-4-2-4-4-1), undulations present ……. ……………………………………………………………………………………………………………………………………………….D. mariae
The zappalai group:
22(3). Dorsal cuticle smooth or slightly rugose …………………………………………………………………………………………………. 23
-. Dorsal cuticle with small tubercles …………………………………………………………………………………………… D. taiwanus
23(22). Oral cavity armature present ………………………………………………………………………………………………………………. 24
-. Oral cavity armature absent/not visible under LM) ………………………………………………………………………………….. 26
24(23). Oral cavity armature composed of a single dorsal tooth ………………………………………………………………………….. 25
-. Oral cavity armature with one anterior band of large teeth and one single medio-dorsal tooth in the posterior band ………………………………………………………………………………………………………………………………………. D. turkmenicus
25(24). Configuration of macroplacoids 1-2-3, secondary branches of claws longer than 50% of the primary branches … …………………………………………………………………………………………………………………………………………. D. mexicanus
-. Configuration of macroplacoids 2-1-3, secondary branches of claws shorter than 50% of the primary branches … …………………………………………………………………………………………………………………………………………. D. bertolanii
26(23). Lunules on claws present, freshwater species …………………………………………………………………………… D. zappalai
-. Lunules on legs absent, terrestrial species ………………………………………………………………………………………………. 27
27(26). Claws without enlarged bases, ratio of the secondary to the primary branches more than 0.79 ……. D. longistipes
-. Claws with enlarged bases, ratio of the secondary to the primary branches less than 0.70 …………………………….. 28
28(27). Ratio of the secondary to the primary branches of claws more than 0.60 …………………………………. D. qinlingense
– Ratio of the secondary to the primary branches of claws less than 0.55 …………………………………… D. korganovae


Citations

Doryphoribius image from Beasley CW, Kaczmarek Ł, Michalczyk Ł. 2008. Doryphoribius mexicanus, a new species of Tardigrada (Eutardigrada: Hypsibiidae) from Mexico (North America). Proceedings of the Biological Society of Washington. 121 (1): 34-40.

“Parts” images 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.

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

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

Schuster RO, Nelson DR, Grigarick AA, Christenberry D. 1980. Systematic criteria of the Eutardigrada. Transactions of the American Microscopical Society. 99: 284-303.

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