RESOLVING THE PARAPHYLETIC TURTLE BLOOD FLUKES: REVISION OF SPIRORCHIIDAE STUNKARD, 1921 AND PROPOSAL OF CARETTACOLIDAE YAMAGUTI, 1958, HAPALOTREMATIDAE (STUNKARD, 1921) POCHE, 1926, BARACKTREMATIDAE N. FAM., PLATTIDAE N. FAM., AND ATAMATAMIDAE N. FAM.

Turtle blood flukes ("spirorchiids") comprise a paraphyletic assemblage including the monophyletic Schistosomatidae Stiles and Hassall, 1898 as a crown group. We herein morphologically diagnose the natural groups of turtle blood flukes and propose family names for them. Spirorchiidae Stunkard, 1921 (Spirorchis MacCallum, 1919 [type]; Spirhapalum Ejsmont, 1927; Plasmiorchis Mehra, 1934; MonticelliusMehra, 1939; VasotremaStunkard, 1928; provisionally UterotremaPlatt and Pichelin, 1994) has a ventral sucker (lost in Spirorchis), an esophageal gland surrounding the entire esophagus, lateral esophageal diverticula (plicate organ and medial esophageal diverticulum present or absent), a glandular mass at the esophagus base, paired ceca surrounded by vitelline follicles for their entire length, a non-filamented, ovoid egg, and typically a Manter's organ. Baracktrematidae n. fam. (Baracktrema Roberts, Platt, and Bullard, 2016 [type]; Unicaecum Stunkard, 1925; NeospirorchisPrice, 1934) has no ventral sucker, a single cecum or cyclocoel, a coiled or looped testis extending into the anterior body half, a convoluted post-cecal and post-gonadal uterus, a uterine pouch, no metraterm, no Manter's organ, and non-filamented eggs. Plattidae n. fam. (Platt Roberts and Bullard, 2018 [type]; Hapalorhynchus Stunkard, 1922; Coeuritrema Mehra, 1933; Enterohaematotrema Mehra, 1940; CardiotremaDwivedi, 1967; Ruavermis Dutton and Bullard, 2020) has the anatomical sequence of a ventral sucker, external seminal vesicle, cirrus sac, anterior testis, ovary, transverse vitelline duct, and posterior testis (except Enterohaematotrema) plus a pars prostatica, dorsal genital pore, globular excretory vesicle, and no Manter's organ. Carettacolidae Yamaguti, 1958 (CarettacolaManter and Larson, 1950 [type]) has a spinose oral sucker, a spinose and contractile ventral sucker (transverse cavity present), numerous testes distributing in a post-ovarian, inter-cecal column, and an ovary and transverse vitelline duct that are pre-testicular. Hapalotrematidae (Stunkard, 1921) Poche, 1926 (Hapalotrema Loos, 1899 [type]; AmphiorchisPrice, 1934; LearediusPrice, 1934; Cheloneotrema Simha and Chattopadhyaya, 1980; NeocaballerotremaSimha, 1977; Satyanarayanotrema Simha and Chattopadhyaya, 1980; Shobanotrema Simha and Chattopadhyaya, 1980) has the anatomical sequence of a ventral sucker, anterior testis(es), external seminal vesicle and cirrus sac, ovary, and posterior testis(es). Atamatamidae n. fam. (Atamatam Bullard and Roberts, 2019 [type]; Paratamatam Bullard and Roberts, 2019; Pitiutrema Dutton and Bullard, 2019) has an oral sucker with anteroventral spines, vasa efferentia connecting to a pre-ovarian external seminal vesicle, a cirrus sac directed postero-laterad and having an internal seminal vesicle and glandular luminal wall, and a dendritic ovary flanked by the external seminal vesicle and oviducal seminal receptacle. A 28S phylogenetic analysis failed to reject the monophyly of these families.