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Figs 10 A–C. Notohymena australis after protargol impregnation. A in Two Oxytrichids from the Ancient Lake Biwa, Japan, with Notes on Morphogenesis of Notohymena australis (Ciliophora, Sporadotrichida)

Figs 10 A–C. Notohymena australis after protargol impregnation. A – venral view of a very late divider showing infraciliature and parental cirri (arrows); B, C – ventral (B) and dorsal (C) view of the proter just after division showing infraciliature and nuclear apparatus, parental cirri (arrows) and newly formed caudal cirri (arrowheads). 1, 5, 6 – dorsal kinety 1, 5, 6, E – endoral membrane, LMR – left marginal row, P – paroral membrane, RMR – right marginal row. Scale bars: 40 μm.

Figs 4A–D. Notohymena australis after protargol impregnation. A in Two Oxytrichids from the Ancient Lake Biwa, Japan, with Notes on Morphogenesis of Notohymena australis (Ciliophora, Sporadotrichida)

Figs 4A–D. Notohymena australis after protargol impregnation. A – ventral view of a representative specimen showing infraciliature and macronuclear nodules (arrows); B – ventral view of cell with two pretransverse ventral cirri (arrows), one additional ventral cirrus (doublearrowhead), and three frontal cirri (arrowheads); C – ventral view of cell, showing infraciliature, macronuclear nodules and micronuclei (arrows), buccal cirrus (arrowhead), and distal end of adoral zone (double-arrowhead); D – anterior part of specimen in dorsal view, showing dorsal kineties. AZM – adoral zone of membranelles, CC – caudal cirri, E – endoral membrane, FVC – frontoventral cirri, 1–6 – dorsal kinety 1–6, LMR – left marginal row, P – paroral membrane, PTVC– pretransverse ventral cirri, PVC– postoral ventral cirri, RMR – right marginal row, TC – transverse cirri. Scale bars: 40 μm.

Figs 9A–H. Notohymena australis after protargol impregnation. A in Two Oxytrichids from the Ancient Lake Biwa, Japan, with Notes on Morphogenesis of Notohymena australis (Ciliophora, Sporadotrichida)

Figs 9A–H. Notohymena australis after protargol impregnation. A – ventral view of middle part of an early divider, showing oral primordium; B, C – ventral view of proter (B) and opisthe (C) of the same middle divider, showing the formation of frontal-ventral-transverse cirral anlagen in proter and opisthe, and anlagen of marginal rows (arrows), and dedifferentiation of anterior end of undulating membranes (arrowhead); D – ventral infraciliature of a proter of a middle divider, showing frontal-ventral-transverse cirral anlagen beginning to organize into new cirri from anterior (arrows) and a small anlage separating from anterior right end of undulating membranes anlagen (arrowhead); E – part of a middle divider in dorsal view, showing fused macronuclear mass (arrow); F – ventral infraciliature of the same divider as in E, showing development of anlagen of the leftmost frontal cirrus (arrowheads), and frontal-ventral-transverse cirral anlagen (arrows); G – ventral infraciliature of a middle-late divider, showing frontal-ventral-transverse cirral anlagen segregating into new cirri (arrows), the anlage of the leftmost frontal cirrus separating from undulating membrane anlagen (arrowheads), and fused macronuclear mass; H – ventral infraciliature of a late divider, showing the formation of anlagen of dorsomarginal kineties at anterior right end of right marginal rows anlagen (arrows) and anterior part of newly formed adoral zone of membranelles curving to right in the opisthe, frontal-ventral-transverse cirral anlagen II – VI segregating into 3, 3, 3, 4 and 4 cirri, the leftmost frontal cirrus originated from undulating membranes anlagen (arrowheads). DKA – dorsal kinety anlage, LMA – anlage of left marginal row, OP – oral primordium, RMA – anlagen of right marginal row. Scale bars: 40 μm.

Figs 6A–D. Notohymena australis after protargol impregnation. A, B in Two Oxytrichids from the Ancient Lake Biwa, Japan, with Notes on Morphogenesis of Notohymena australis (Ciliophora, Sporadotrichida)

Figs 6A–D. Notohymena australis after protargol impregnation. A, B – ventral (A) and dorsal (B) infraciliature of the same very late divider, showing parental cirri (arrows) and newly formed caudal cirri at posterior ends of first, second and fourth dorsal kineties (arrowheads); C, D – ventral (C) and dorsal (D) infraciliature of the proter just after division showing two macronuclear nodules, one micronucleus, parental cirri (arrows), and newly formed caudal cirri at the posterior ends of first, second and fourth dorsal kineties (arrowheads). 1–6 – dorsal kinety 1–6, LMR – left marginal row, Ma – macronuclear nodule, Mi – micronucleus, RMR – right marginal row. Scale bars: 40 μm.

Figs 8A–M. Notohymena australis after protargol impregnation. A in Two Oxytrichids from the Ancient Lake Biwa, Japan, with Notes on Morphogenesis of Notohymena australis (Ciliophora, Sporadotrichida)

Figs 8A–M. Notohymena australis after protargol impregnation. A – ventral view of a representative specimen showing infraciliature; B – plumper cell showing pharyngeal fibers (arrowhead), macronuclear nodules, and micronuclei (arrows); C, L – ventral infraciliature of the same cell with additional ventral cirrus (arrow); D –ventral view of middle part of a specimen with 4 postoral ventral cirri; E – anterior part of the same cell as in A, showing hooked anterior end of paroral membrane (arrow) and three enlarged frontal cirri (arrowheads); F, G – the same individual showing ventral ciliature and ingested ciliate (arrow, Aspidisca sp.); H – details of buccal apparatus showing endoral membrane and the structure of adoral membranelles; I – macronuclear nodules composed of many small nucleoi and micronuclei (arrows); J, M – ventral view of anterior part of cell, showing cirri associated with fibers; K – ventral view of posterior part of cell, showing left and right marginal rows terminating respectively at posterior end and at level of right transverse cirrus (arrows), and caudal cirri (arrowheads). AZM – adoral zone of membranelles; E – endoral membrane, LMR – left marginal row, Ma – macronuclear nodule, P – paroral membrane, RMR – right marginal row, TC – transverse cirri. Scale bars: 40 μm.

Figs 3A–Q. Apoamphisiella tihanyiensis after protargol impregnation. A in Two Oxytrichids from the Ancient Lake Biwa, Japan, with Notes on Morphogenesis of Notohymena australis (Ciliophora, Sporadotrichida)

Figs 3A–Q. Apoamphisiella tihanyiensis after protargol impregnation. A – ventral view of specimen with two frontal cirri, showing micronuclei (arrows) and caudal cirri (arrowhead); B – right lateral view showing several parental cirri (arrow); C – ventral view of a specimen with two additional ventral cirri ahead of transverse cirri; D, F – ventral view of anterior part of an individual with two frontoventral cirri; E, G – ventral view of anterior part of a cell with three frontoventral and two postoral ventral cirri (arrowheads); H – ventral view of cell with several parental cirri (arrow), and single postoral ventral cirrus (arrowhead); I – ventral view of middle part of cell, showing oral primordium; J – ventral view of a cell just after division, showing anteriorly placed postoral ventral cirrus (arrow); K – dorsal view of a cell, showing dorsal kineties and macronuclear nodules; L – ventral view of posterior part of specimen with four transverse cirri; M – ventral view of middle part of cell, showing cirri associated with fibres; N – ventral view of middle part of a cell, showing single postoral ventral cirrus (arrow) and parental cirri (arrowhead); O – dorsal view of anterior part of a cell, showing dorsal kineties and associated cortical granules; P, Q – ventral views of posterior part of cell, showing caudal cirri (arrows). DK – dorsal kinety, E – endoral membrane, FC – frontal cirri, Ma – macronuclear nodule, Mi – micronucleus, P – paroral membrane, TC – transverse cirri. Scale bars: 40 μm.

Figs 2A–H. Apoamphisiella tihanyiensis from life. A, B in Two Oxytrichids from the Ancient Lake Biwa, Japan, with Notes on Morphogenesis of Notohymena australis (Ciliophora, Sporadotrichida)

Figs 2A–H. Apoamphisiella tihanyiensis from life. A, B – ventral view of different cells; C – a contracted cell, showing contractile vacuole; D – part of cell in dorsal view, showing contractile vacuole with collecting canals (arrow); E, F – ventral views of anterior (E) and posterior (F) parts of the same cell, showing arrangement of cortical granules; G – plumper individual, showing cortical granules (arrowheads) and macronuclear nodules; H – part of cell in dorsal view showing spherical contractile vacuole and arrangement of cortical granules. CV – contractile vacuole, FC – frontal cirri, Ma – macronuclear nodule, RMR – right marginal row, TC – transverse cirri. Scale bars: 60 μm.

Figs 1A–L in Two Oxytrichids from the Ancient Lake Biwa, Japan, with Notes on Morphogenesis of Notohymena australis (Ciliophora, Sporadotrichida)

Figs 1A–L. Apoamphisiella tihanyiensis from life (A–D) and after protargol impregnation (E–L). A – ventral view of a representative individual; B – arrangement of cortical granules on ventral side; C – shape of contractile vacuole; D – arrangement of cortical granules on dorsal side; E – posterior part of cell in ventral view, showing 2 pretransverse ventral cirri (arrowheads) and two additional ventral cirri ahead of transverse cirri (arrows); F – posterior part of cell in ventral view, showing 2 pretransverse ventral cirri (encircled by broken line) and caudal cirri (arrows); G, H – ventral (G) and dorsal (H) infraciliature of the same specimen, showing single postoral ventral cirrus (arrow in G), caudal cirri at the posterior end of dorsal kinety 4 (arrow in H), and frontal cirri (arrowheads in G); I – anterior half of cell in ventral view, showing single postoral ventral cirrus (arrow) and several parental ventral cirri (arrowhead); J – part of ventral infraciliature, showing micronuclei (arrows), frontoventral cirri (arrowheads), 2 postoral ventral cirri (double-arrowhead), and several parental ventral cirri (connected by broken line); K, L – ventral (K) and dorsal (L) infraciliature of the same specimen, showing caudal cirri (arrows in K), macronuclear nodules (arrows in L), frontal cirri (arrowheads in K), and micronuclei (arrowheads in L). 1–6 – dorsal kinety 1–6, AZM, adoral zone of membranelles; BC – buccal cirrus, CR – cirral row, DB – dorsal bristle, E – endoral membrane, FVC – frontoventral cirri, FVR – frontoventral row, LMR – left marginal row, Ma – macronuclear nodule, P – paroral membrane, RMR – right marginal row, TC – transverse cirri. Scale bars: 80 μm (A), 40 μm (I, G, H, J–L), 20 μm (E, F).

Figs 7A–I. Notohymena australis from life. A, G in Two Oxytrichids from the Ancient Lake Biwa, Japan, with Notes on Morphogenesis of Notohymena australis (Ciliophora, Sporadotrichida)

Figs 7A–I. Notohymena australis from life. A, G – ventral view of the same cell, showing flexible body, contractile vacuole, ingested algae (arrow) and macronuclear nodules; B, C, D – ventral view of individuals in different states, showing hooked anterior end of peristomial lip (arrow); E, I – view of portion of cell, showing the arrangement of yellow-greenish cortical granules, and dorsal cilia (arrows in E and arrowheads in I); F – macronuclear nodules, micronuclei (arrows) and cortical granules (arrowheads); H – dorsal view of a curved cell, showing cortical granules grouped in lines. CV – contractile vacuole, Ma – macronuclear nodule, TC – transverse cirri. Scale bars: 40 μm.

Figs 5A–G. Notohymena australis after protargol impregnation. A in Two Oxytrichids from the Ancient Lake Biwa, Japan, with Notes on Morphogenesis of Notohymena australis (Ciliophora, Sporadotrichida)

Figs 5A–G. Notohymena australis after protargol impregnation. A – ventral view of an early divider, showing oral primordium; B, C – ventral (B) and dorsal (C) view of the same middle divider, showing the formation of frontal-ventral-transverse cirral anlagen in proter and opisthe, and anlagen of marginal rows (arrows), dedifferentiation of anterior end of undulating membranes (arrowheads), and undulating membrane anlagen in opisthe (double-arrowhead); D, E – ventral infraciliature of two middle dividers, showing frontal-ventral-transverse cirral anlagen beginning to organize into new cirri from anterior (arrowheads) and a small anlage separating from anterior right end of undulating membrane anlagen (arrows), insets: fused macronucleus; F, G – ventral (B) and dorsal (C) view of the same late divider, showing the formation of anlgen of dorsomarginal kineties at anterior right end of right marginal rows anlagen (arrows in F), frontal-ventral-transverse cirral anlagen II – VI segregating into 3, 3, 3, 4 and 4 cirri, the leftmost frontal cirrus originated from undulating membranes anlagen (arrowheads), and development of dorsal kinety anlagen (arrows in G). 1–4 – dorsal kinety 1–4, LMA – anlage of left marginal row, OP – oral primordium, RMA – anlagen of right marginal row. Scale bars: 40 μm.

FIGURE 11 in Diversification in ancient Lake Biwa: integrative taxonomy reveals overlooked species diversity of the Japanese freshwater snail genus Semisulcospira (Mollusca: Semisulcospiridae)

FIGURE 11 Shells of Semisulcospira salebrosa sp. nov. A–G, Holotype, KUZ Z4131. H–J, Paratype, KUZ Z4133. K–L, Paratype, KUZ Z4135. M–O, Specimen from Take-shima Island, KUZ Z4138. A–C, H, K, M, Adult shell. A–C, H, M, Female. K, Male. D, I, L, N, Operculum. E–G, J, O, Embryonic shell. Scale bars: 10 mm (A–D, H–I, K–L, M–N), 1 mm (E–G, J, O). All specimens were treated with 3% sodium hypochlorite

FIGURE 9 in Diversification in ancient Lake Biwa: integrative taxonomy reveals overlooked species diversity of the Japanese freshwater snail genus Semisulcospira (Mollusca: Semisulcospiridae)

FIGURE 9 Shells of Semisulcospira watanabei sp. nov. A–G, Holotype, KUZ Z4109. H–J, Paratype, KUZ Z4110. K–L, Paratype, KUZ Z4114. M–N. Specimen from Oura, KUZ Z4117. O–Q, Specimen from Nihonmatsu, KUZ Z4118. R–T, Specimen from Horikiri Port, KUZ Z4120. A–C, H, K, M, O, R, Adult shell. A–C, H, O, R, Female. K, Male. M, Juvenile. D, I, L, N, P, S, Operculum. E–G, J, Q, T, Embryonic shell. Scale bars: 10 mm (A–D, H–I, K–L, M–N, O–P, R–S), 1 mm (E–G, J, Q, T). All specimens were treated with 3% sodium hypochlorite

FIGURE 7 in Diversification in ancient Lake Biwa: integrative taxonomy reveals overlooked species diversity of the Japanese freshwater snail genus Semisulcospira (Mollusca: Semisulcospiridae)

FIGURE 7 Results of Random Forest analyses conducted for five Semisulcospira species. Euclidean distances generated from proximities among individuals are plotted. A, Female. B, Male

FIGURE 6 in Diversification in ancient Lake Biwa: integrative taxonomy reveals overlooked species diversity of the Japanese freshwater snail genus Semisulcospira (Mollusca: Semisulcospiridae)

FIGURE 6 Map of Lake Biwa indicating geographical variation in the frequency of sculpture types in the adult females of five Semisulcospira species. Colours of locality names correspond to the colour coding in fig. 1

FIGURE 4 in Diversification in ancient Lake Biwa: integrative taxonomy reveals overlooked species diversity of the Japanese freshwater snail genus Semisulcospira (Mollusca: Semisulcospiridae)

FIGURE 4 Results of ADMIXTURE analysis based on 738 SNP s conducted for five Semisulcospira species. Bar colours of the species in K = 5 correspond to the colour coding in fig. 3

FIGURE 5 in Diversification in ancient Lake Biwa: integrative taxonomy reveals overlooked species diversity of the Japanese freshwater snail genus Semisulcospira (Mollusca: Semisulcospiridae)

FIGURE 5 Phylogenetic relationships of five Semisulcospira species estimated by the Neighbor-Net reconstructed based on uncorrected p-distances of 738 SNP s

FIGURE 3 in Diversification in ancient Lake Biwa: integrative taxonomy reveals overlooked species diversity of the Japanese freshwater snail genus Semisulcospira (Mollusca: Semisulcospiridae)

FIGURE 3 Results of principal components analysis based on 738 SNP s conducted for five Semisulcospira species. A, Principal component (PC) 1 vs PC 2. B, PC2 vs PC3. C, PC 3 vs PC4. D, PC4 vs PC5

FIGURE 2 in Diversification in ancient Lake Biwa: integrative taxonomy reveals overlooked species diversity of the Japanese freshwater snail genus Semisulcospira (Mollusca: Semisulcospiridae)

FIGURE 2 Schematic drawings representing shell measurements of Semisulcospira species in this study. A, Spire angle (SA) and aperture swell length (ASL) of adult shell. B–D, Criteria for types of sculptures

FIGURE 1 in Diversification in ancient Lake Biwa: integrative taxonomy reveals overlooked species diversity of the Japanese freshwater snail genus Semisulcospira (Mollusca: Semisulcospiridae)

FIGURE 1 Map of Lake Biwa (A) and Oura Bay (B) showing 15 sampling localities: blue, Semisulcospira niponica; green, S. fuscata; purple, S. watanabei sp. nov.; red, S. nakanoi sp. nov.; orange, S. salebrosa sp. nov; black, putative hybrid between S. fuscata and S. watanabei sp. nov.

FIGURE 10 in Diversification in ancient Lake Biwa: integrative taxonomy reveals overlooked species diversity of the Japanese freshwater snail genus Semisulcospira (Mollusca: Semisulcospiridae)

FIGURE 10 Shells of Semisulcospira nakanoi sp. nov. A–G, Holotype, KUZ Z4122. H–J, Paratype, KUZ Z4125. K–L, Paratype, KUZ Z4126. M–Q, Specimens from Onoe Port, KUZ Z4129, Z4130. A–C, H, K, M, P, Adult shell. A–C, H, M, Female. K, P, Male. D, I, L, N, Q, Operculum. E–G, J, O, Embryonic shell. Scale bars: 10 mm (A–D, H–I, K–L, M–N, P–Q), 1 mm (E–G, J, O). All specimens were treated with 3% sodium hypochlorite