Haloklossiinae, Tokiwa, Chou, 2021

Tokiwa, Toshihiro, Chou, Shyun, Tochigi, Yuki, Katayama, Kentaro & Duszynski, Donald W., 2021, Hyaloklossia Labb´e, 1896 (Alveolata: Apicomplexa) in frogs: Description of a new species and proposing a new subfamily to accommodate these enigmatic parasites, International Journal for Parasitology: Parasites and Wildlife 15, pp. 199-207 : 205

Family Sarcocystinae Poche, 1913

Subfamily Haloklossiinae n. subfam. Tokiwa, Chou, Tochigi, Katayama, et Duszynski (2021).

Diagnosis: Members exhibit a homoxenous life cycle and undergo endogenous sporulation in the kidneys of anurans. Oocysts with two sporocysts each with four sporozoites. Stieda and substieda bodies absent. Sporocyst residuum present.

Type genus: Hyaloklossia Labb´e, 1896

with high bootstrap values [18S: 95 (NJB)/91 (MLB); 28S: 100 (NJB)/ 100 (MLB); cox1: 100 (NJB)/100 (MLB)], whereas clade Ib composed of Besnoitia spp. had moderate to high bootstrap values [18S: 70 (NJB)/72 (MLB); 28S: 97 (NJB)/88 (MLB)]. The Eumonosporinae clade (clade II) was monophyletic and a sister group of the Toxoplasmatinae (clade Ia + Ib) with low to moderate bootstrap values [18S: 55 (NJB)/51 (MLB); 28S: 58 (NJB)/64 (MLB); cox1: 57 (NJB)/61 (MLB)]. Hyaloklossia spp . (clade III) and Cystoisosporinae (clade IV) were monophyletic groups, respectively. With respect to the 18S and cox1 sequences, these groups collectively formed a monophyletic clade (III + IV) with moderate bootstrap values (above 50%), whereas with respect to the 28S sequence, clade III showed the formation of an early branch at the monophyletic clade comprising Ia + Ib + II + IV. The phylogenetic positions of Nephroisospora were unstable, and it was placed in clade I (based on 18S sequence) or clade IV (based on cox1 sequence).