Doryphoribiidae, Gąsiorek & Stec & Morek & Michalczyk, 2019

Gąsiorek, Piotr, Stec, Daniel, Morek, Witold & Michalczyk, Łukasz, 2019, Deceptive conservatism of claws: distinct phyletic lineages concealed within Isohypsibioidea (Eutardigrada) revealed by molecular and morphological evidence, Contributions to Zoology 88 (1), pp. 78-132 : 130-132

Composition of Doryphoribiidae fam. nov.

al., 1980, D. neglectus Pilato & Lisi, 2004 , D. pilatoi Bertolani, 1984 , D. smokiensis Bartels et al., 2007 ;

– two macroplacoids present, dorsolateral gibbosities absent, aquatic (the D. koreanus group): D. koreanus Moon et al., 1994 , D. polynettae Biserov, 1988 , D. tergumrudis Bartels et al., 2008 ;

Genus: Apodibius Dastych, 1983 – three macroplacoids and dorsolateral gibbosities present, terrestrial (the D. vietnam-

Species: A. confusus Dastych, 1983 , A. nuntius ensis group): D. elleneddiei Haefƙe et al., 2014 , Binda, 1984, A. richardi Vargha, 1995 D. gibber Beasley & Pilato, 1987 , D. maasaima-

rensis Fontoura et al., 2013, D. mariae Pilato

Genus: Doryphoribius Pilato, 1969 & Binda, 1990, D. minimus Bartels et al., 2008 ,

D. vietnamensis (Iharos, 1969) ;

Species (divided into ecomorphological groups distinguished in this paper): – three macroplacoids present, dorsolateral gibbosities absent, terrestrial (the D. bertola-

– two macroplacoids and dorsolateral gibbos- nii group): D. bertolanii Beasley & Pilato, 1987 , ities present, terrestrial (the D. flavus group): D. chetumalensis Pérez-Pach et al., 2017 , D. kor- D. amazzonicus Lisi, 2011 , D. barbarae Beasley ganovae Biserov, 1994, D. mexicanus Beasley et & Miller, 2012, D. bindae Lisi, 2011 , D. dawkinsi al., 2008, D. qinlingense Li et al., 2004 , D.taiwa- Michalczyk & Kaczmarek, 2010, D. dupli- nus Li & Li, 2008, D. turkmenicus Biserov, 1999 ; globulatus Ito, 1995, D. flavus (Iharos, 1966) , D. huangguoshuensis Wang et al., 2007 , D. ma- – three macroplacoids present, dorsolatranguensis Binda & Pilato, 1995, D. mcinnesae eral gibbosities absent, aquatic (the D. zap- Meng et al., 2014, D. monstruosus (Maucci, palai group): D. longistipes Bartels et al., 2008 , 1991) comb. nov., D. niedbalai Zawierucha D. zappalai Pilato, 1971 . et al., 2012, D. picoensis Fontoura et al., 2008 , D. quadrituberculatus Kaczmarek & Michalc- Remarks: Currently, Doryphoribius comprises zyk, 2004, D. rosanae Daza et al., 2017 , D. sol- the most diverse group of taxa in terms of claw idunguis Lisi, 2011, D. tessellatus Meyer, 2011 , morphology within the entire Isohypsibioidea D. zyxiglobus (Horning et al., 1978) ; (see Discussion),which suggests that the genus is polyphyletic (Bertolani et al., 2014a; Cesari et

– two macroplacoids and dorsolateral gibbos- al., 2016;Gąsiorek et al.,2019).In the light of our ities present, aquatic (the D. evelinae group): findings regarding the polyphyly of Isohypsibi- D. evelinae (Marcus, 1928) ; us, it seems very likely that Doryphoribius may comprise a number of new genera.

– two macroplacoids present, dorsolateral Given that Isohypsibius monstruosus Maucgibbosities absent, terrestrial (the D. dorypho- ci, 1991 exhibits the ventral lamina ( fig. 17A ), rus group = Doryphoribius s.s.): D. doryphorus which conforms with the current diagnosis (Binda & Pilato, 1969), D. macrodon Binda et of Doryphoribius Downloaded , the from new Brill.com combination 12/12/2023 02 D:59.:51PM

via Open Access. This is an open access article distributed under the terms of the prevailing CC-BY license at the time of publication.

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monstruosus (Maucci, 1991) comb. nov. is pro- Remarks: All former aquatic Isohypsibius speposed. The transfer confirms the supposition cies are ascribed to the new genus. Although of Michalczyk & Kaczmarek (2010), that some Grevenius gen. nov. comprises taxa dwelling Isohypsibius spp. may in fact represent Dory- in similar habitats, clear peculiarities can be phoribius s.l. observed in claw morphology of some limnic and intertidal species ( G.brevitubulatus comb. Genus: Grevenius gen. nov. nov., G. deflexus comb. nov., G. granditintinus comb. nov., G. hydrogogianus comb. nov., Species: G. annulatus annulatus (Murray, G.irregibilis comb. nov., G.myrops comb. nov.). 1905) comb. nov., G. annulatus minor (Ramaz- Considering that even slight differences in zotti, 1945) comb. nov., G. asper (Murray, 1906) claw anatomy seem to hold very strong phylocomb. nov., G. baicalensis (Ramazzotti, 1966) genetic signal in Isohypsibioidea , it will not be comb. nov., G. baldii (Ramazzotti, 1945) comb. surprising if these species turn out to represent nov., G. baldiioides (Tumanov, 2003) comb. separate genera when more accurate morphonov., G. brevitubulatus (Rho et al., 1997) comb. logical data and DNA sequences are available. nov., G. deconincki (Pilato, 1971) comb. nov., G. karenae comb. nov., G. rugosus comb. G.deflexus (Mihelčič,1960) comb.nov., G.fuscus nov. and G. sismicus comb. nov. were found (Mihelčič, 1971/72) comb. nov. et nom. inq., in hydrophilic substrata, in close vicinities of G. granditintinus (Chang & Rho, 1996) comb. lakes or ponds, but not strictly in an aquatic nov., G. granulifer (Thulin, 1928) comb. nov., habitat. However, their close affinity to Greve- G. hydrogogianus (Ito & Tagami, 1993) comb. nius gen. nov. seems certain, especially that nov., G. irregibilis (Biserov, 1992) comb. nov., the most similar species to which they were G. karenae (Zawierucha, 2013) comb. nov., compared in their original descriptions were G. kenodontis (Kendall-Fite & Nelson, 1996) all exclusively limnic taxa. comb. nov., G. koreanensis (Iharos, 1971) comb. Due to morphological differences with nov. et nom. inq., G. kotovae (Tumanov, 2003) G. granulifer comb. nov. (pink body colour comb. nov., G. kristenseni (Pilato et al., 1989) and cuticular tubercles of identical size on comb. nov., G. ladogensis (Tumanov, 2003) the whole body in Isohypsibius granulifer kocomb . nov., G. laevis (McInnes, 1995) comb. reanensis vs white body colour and cuticular nov., G. lineatus (Mihelčič, 1969) comb. nov. et tubercles of different sizes on the dorsum nom. inq., G. longiunguis (Pilato, 1974) comb. and on the venter in G. granulifer comb. nov.), nov., G. malawiensis (Jørgensen, 2001) comb. I. granulifer koreanensis is elevated to a spenov., G. marii (Bertolani, 1981) comb. nov., cies rank as Grevenius koreanensis comb. nov. G. monoicus (Bertolani,1981) comb. nov., G.my - rops (du Bois-Reymond Marcus, 1944) comb. Genus: Paradiphascon Dastych, 1992 nov., G.nipponicus (Sudzuki,1975) comb.nov. et nom. inq., G. pulcher (Mihelčič, 1971/72) comb. Species: P. manningi Dastych, 1992 . nov. et nom. inq., G. pushkini (Tumanov, 2003) comb. nov., G. rugosus (Guidi & Grabowski, Remarks:Given the peculiar apomorphy in the 1996) comb. nov. et nom. inq., G. rusticus (Pila- form of flexible pharyngeal tube and dispro - to et al., 2015) comb. nov., G. sismicus (Maucci, portionally widened bases of external and pos- 1978) comb. nov., G. tubereticulatus (Pilato & terior claws, the genus requires an integrative Catanzaro, 1989) comb. nov., G. verae (Pilato redescription to verify its systematic position. & Catanzaro, 1989) comb. nov., G. zappalai Downloaded from Brill.com 12/12/2023 02:59:51PM (Pilato et al., 2015) comb. nov. of via Open Access. This Genus the is prevailing an: Pseudobiotus open access CC-BY license article Nelson at the, 1980 distributed under the terms time of publication. http://creativecommons.org/licenses/by-nc/4.0

Species: P. hirsutellus Pilato et al., 2010 , P.kathmanae Nelson et al., 1999, P. longiunguis (Iharos, 1968) sp. dub., P. matici (Pilato, 1971) , P. megalonyx (Thulin, 1928) , P. spinifer Chang et al., 2007 , P. vladimiri Biserov et al., 2001 .

Remarks: The oldest species in the genus, P.megalonyx ,requires an integrative redescription to aid species discovery in Pseudobiotus .

Genus: Thulinius Bertolani, 2003

Species: T. augusti (Murray, 1907) , T. itoi (Tsurusaki, 1980) , T. romanoi Bertolani et al., 2014 , T.ruffoi (Bertolani, 1981) , T. saltursus (Schuster et al., 1978) , T. stephaniae (Pilato, 1974) .

Remarks: The oldest species in the genus, T. augusti , requires an integrative redescription to allow for the verification of alleged numerous records of the species throughout the world and to aid species discovery in Thulinius .

Figures

figure 17 Doryphoribius monstruosus (Maucci, 1991) comb. nov. (PCM, holotype): A – the buccal apparatus (arrowhead indicates ventral lamina); B – claws I; C – claws IV. Scale bars in micrometres