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Figure 7 in Morphological, ontogenetic, and molecular investigations of freshwater hypotrich ciliates from China revealed a new genus Heterodeviata gen. nov. (Protista: Ciliophora), and a novel limnetic population of Deviata multilineae
Figure 7. Deviata multilineae from life (A–I) and asser protargol impregnation (J–N). A, F, ventral view of a representative individual, arrowhead in (F) marks the algae in the cell. B–D, G, H, ventral views of different cells, arrowheads indicate contractile vacuole. E, ventral view, showing the detail of cirri and their surrounded globules. I, denoting the macronuclear nodules and the cytoplasm. J, K, N, ventral (J, N) and dorsal (K) view of the same representative cell, showing infraciliature and nuclear apparatus, arrowhead in (J) marks the buccal cirrus, arrows and arrowheads in (K, N) indicate macronuclear nodules and micronuclei, respectively. L, dorsal view to show the less marginal rows on dorsal side. M, ventral view, arrowhead indicates buccal cirrus. 1–3, dorsal kineties 1–3; E, endoral; FC, frontal cirri; FVR1–3, frontoventral cirral rows 1–3; LMR, less marginal row; Ma, macronuclear nodules; P, paroral; PBC, parabuccal cirri; RMR, right marginal row. Scale bars: 50 μm.
Figure 8 in Morphological, ontogenetic, and molecular investigations of freshwater hypotrich ciliates from China revealed a new genus Heterodeviata gen. nov. (Protista: Ciliophora), and a novel limnetic population of Deviata multilineae
Figure 8. Nucleotide differences between deviatids based on 18S rRNA gene sequences. In (A), the lower less values of the table indicate the sequence similarity, the upper right numbers are numbers of nucleotide differences. The numbers in the header (B) indicate the unmatched site positions. In A, B, the newly obtained sequences are in red. '**' indicates the sequence Perisincirra sp. (KY855575) which was likely misidentified and can be considered conspecific with Deviata brasiliensis. 'H', Heterodeviata.
Figure 5 in Morphological, ontogenetic, and molecular investigations of freshwater hypotrich ciliates from China revealed a new genus Heterodeviata gen. nov. (Protista: Ciliophora), and a novel limnetic population of Deviata multilineae
Figure 5. Morphogenetic process of Heterodeviata sinica asser protargol impregnation. A, showing the oral primordium of the opisthe. B, ventral view of a very early divider, showing oral primordium of the opisthe (arrowhead). C–E, ventro-lateral (C, D) and dorsolateral (E) views of the same early divider, in this stage, anlagen I–IV (C), two right marginal anlagen (E) formed, adoral zone of membranelles incompletely formed (arrowhead in C), and arrowhead in (D) shows the parental endoral beginning to disintegrate. F, G, ventral (F) and dorsal (G) views of two middle dividers, arrowheads in (G) showing dorsal kinety anlage 2. H, I, ventral (H) and dorsal view (I) of two middle dividers, in this stage, anlagen I–IV dedifferentiating into cirri (H), and two bristles of dorsal kinety 2 formed (arrowheads in H, I). J, dorsal view of a late divider, showing one macronuclear nodule dedifferentiated into two ones. K, L, ventral views of two late dividers, arrowheads in (K) and (L) indicate the buccal and parabuccal cirrus, separately. I–IV, anlagen I–IV; DK1, dorsal kinety 1; DḎ1, dorsal kinety anlage 1; FVR, frontoventral cirral row; LMA1, 2, less marginal anlagen 1, 2; LMR1, 2, less marginal rows 1, 2; Ma, macronuclear nodules; Mi, micronuclei; OP, oral primordium; RMA1, 2, right marginal anlagen 1, 2; RMR1, 2, right marginal rows 1, 2; UM, undulating membranes. Scale bars: 50 μm (F, L), 20 μm (G, I).
Figure 1 in Morphological, ontogenetic, and molecular investigations of freshwater hypotrich ciliates from China revealed a new genus Heterodeviata gen. nov. (Protista: Ciliophora), and a novel limnetic population of Deviata multilineae
Figure 1. Location of the sampling site. A, map showing the location of the Lake Weishan Wetland, China. B, C, photographs of the Lake Weishan where Heterodeviata sinica and Deviata multilineae were collected, respectively.
Figure 9 in Morphological, ontogenetic, and molecular investigations of freshwater hypotrich ciliates from China revealed a new genus Heterodeviata gen. nov. (Protista: Ciliophora), and a novel limnetic population of Deviata multilineae
Figure 9. Comparison of Heterodeviata with related genera within Deviatidae and Kahllidae that possess three frontal cirri, at least one short or long frontal ventral cirral row right of the cell midline, at least one less marginal row, and one dorsomarginal kinety. Figures in the upper and lower row show the dorsal kinety paưern and cirral paưern of corresponding genera, respectively. Old (parental) structures (marginal cirri or dorsal kineties) are depicted by contour. Abbreviations: BC, buccal cirri; CC, caudal cirri; DK, dorsal kinety; FC, frontal cirri; FVR, frontoventral cirral row; LMR, less marginal row; PBC, parabuccal cirri; RMR, right marginal row; TC, transverse cirri.
Figure 4 in Morphological, ontogenetic, and molecular investigations of freshwater hypotrich ciliates from China revealed a new genus Heterodeviata gen. nov. (Protista: Ciliophora), and a novel limnetic population of Deviata multilineae
Figure 4. Early to middle morphogenetic stages of Heterodeviata sinica asser protargol impregnation. A, macronuclear nodules of a very early divider (ventral view), showing replication band of macronuclear nodules (arrows). B, oral primordium of the opisthe (arrow). C, developed oral primordia (arrow). D–F, ventrolateral (D, E) and dorsolateral (F) view of the same early divider, showing incomplete adoral zone of membranelles (arrow in D, E), the intrakinetally formed marginal anlagen and dorsal kinety anlage 1. G, H, ventral (G) and dorsal (H) view of the same early-middle divider, showing the dorsal kinety 2 anlage formed to the right of the right marginal anlage 2. I, J, ventral (I) and dorsal (J) view of the same middle divider, denoting anlagen I–IV differentiating into cirri, arrowhead in (I) shows the parabuccal cirrus. DḎ1, 2, dorsal kineties anlagen 1, 2; FC, frontal cirri; FVR, frontoventral cirral row; LMA1, 2, less marginal anlagen 1, 2; Ma, macronuclear nodules; Mi, micronuclei; RMA1, 2, right marginal anlagen 1, 2; UMA, undulating membrane anlage. Scale bars: 20 μm (A–C), 50 μm (D–J).
Figure 3 in Morphological, ontogenetic, and molecular investigations of freshwater hypotrich ciliates from China revealed a new genus Heterodeviata gen. nov. (Protista: Ciliophora), and a novel limnetic population of Deviata multilineae
Figure 3. Heterodeviata sinica from life (A, D–F, I–K) and asser protargol impregnation (B, C, G, H, L–P). A, ventral view of a composite representative specimen. B, L, details of the oral zone, arrow in (B) indicates pharyngeal fibres. C, detail of the end of cell, arrowhead indicates the caudal cirrus at the end of dorsal kinety 1. D–F, ventral views, showing the various cell shapes, arrows mark the contractile vacuole. G, H, M–O, ventral (G, N) and dorsal (H, M, O) views of the holotype, denoting infraciliature and nuclear apparatus, arrowhead in (G) marks buccal cirrus, arrowheads in (H, O) and arrow in (M, O) show the two dorsal bristles of dorsal kinety 2 and the single caudal cirrus, respectively. I, J, ventral views to show different individuals, arrow in (I) shows contractile vacuole. K, showing the cytoplasm and macronuclear nodules. P, dorsal view of a different individual, arrowheads and arrow indicate two dorsal bristles and caudal cirrus, respectively. 1, 2, dorsal kineties 1, 2; E, endoral; FC, frontal cirri; FVR, frontoventral cirral row; LMR1, 2, less marginal rows 1, 2; Ma, macronuclear nodules; Mi, micronuclei; P, paroral; PBC, parabuccal cirri; RMR1, 2, right marginal rows 1, 2. Scale bars: 50 μm (A, D–F, I, J), 30 μm (G, H, K–P).
Figure 6 in Morphological, ontogenetic, and molecular investigations of freshwater hypotrich ciliates from China revealed a new genus Heterodeviata gen. nov. (Protista: Ciliophora), and a novel limnetic population of Deviata multilineae
Figure 6. Middle to late morphogenetic stages of Heterodeviata sinica asser protargol impregnation. A, B, ventral (A) and dorsal (B) view of the same middle divider, to demonstrate the formed marginal rows and dorsal kineties 1, 2. C, D, ventral (C) and dorsal (D) view of the same late-middle divider, arrowheads in (C) show the parabuccal cirrus. E, F, ventral (E) and dorsal (F) view of the same late divider, arrowheads in (E) and arrows in (F) separately mark the parabuccal and caudal cirrus for the proter and opisthe. G, H, ventral (G) and dorsal (H) view of the same late divider, arrowheads and arrows separately show the parabuccal and caudal cirrus. BC, buccal cirrus, DK1, 2, dorsal kineties 1, 2; FC, frontal cirri; FVR, frontoventral cirral row; LMR1, 2, less marginal rows 1, 2; Ma, macronuclear nodules; RMR1, 2, right marginal rows 1, 2. Scale bars: 50 μm.
Figure 2. A, maximum likelihood tree inferred from 18S in Morphological, ontogenetic, and molecular investigations of freshwater hypotrich ciliates from China revealed a new genus Heterodeviata gen. nov. (Protista: Ciliophora), and a novel limnetic population of Deviata multilineae
Figure 2. A, maximum likelihood tree inferred from 18S rRNA gene sequences, showing the phylogenetic positions of the two newly sequenced species. Numbers near the nodes represent the ML bootstrap support and BI posterior probability values. Fully supported (100%/1.00) branches are marked with solid circles. 'Asterisks' indicate disagreement between the ML and BI trees. Sequences newly obtained are in pink. The scale bar corresponds to one substitution per 100 nucleotide positions. B, the tree is made referring to the ML tree in a radiation view, showing the likely systematic relationship of Heterodeviata with related genera. C, topology of species within Deviatidae in Bayesian inference (BI) tree. '**' indicates the sequence Perisincirra sp. (KY855575) is probably misidentified, which should be conspecific with Deviata brasiliensis.
Figure 11 in The search finds an end: the morphologically chimeric hysterocinetids belong to the subclass Hymenostomatia (Ciliophora: Oligohymenophorea)
Figure 11. Secondary structure of the 18S rRNA molecule of Hysterocineta bellerophon. Arrows mark molecular synapomorphies of the 'paravorax' clade. Primary nucleotide homologies are scaưered throughout the whole 18S rRNA molecule, suggesting a long common evolution of the 'paravorax' clade. Thus, 'paravorax' clade-specific nucleotide characters are situated in terminal loops of helices (six nucleotide positions), bulges (six positions), single-stranded regions (four positions), and double-stranded regions (32 positions). There are as many as 48 molecular synapomorphies and four indels in the 18S rRNA molecule, corroborating the common origin of hysterocinetids and other members of the 'paravorax' clade. Mutations in the double-stranded regions are typically involved in compensatory base changes or retain helical structure when involved in non-canonical pairings in helices 27 and 43. The 18S secondary structure map of Saccharomyces cerevisiae (inset) is from hưp://apollo.chemistry.gatech.edu/RibosomeGallery.
Figure 8 in The search finds an end: the morphologically chimeric hysterocinetids belong to the subclass Hymenostomatia (Ciliophora: Oligohymenophorea)
Figure 8. Phylogenetic tree based on the nuclear 18S rRNA gene and mitochondrial COI sequences, showing the phylogenetic position of hysterocinetids within the subclass Hymenostomatia. Ichthyophthirius multifiliis was used as an outgroup, following Zhang and Vďačný (2022). Posterior probabilities for Bayesian inference conducted in MrBayes and bootstrap values for maximum likelihood conducted in IQ-Tree were mapped onto the 50%-majority rule Bayesian consensus tree. Fully statistically supported nodes are marked with red solid circles. The scale bar denotes two substitutions per 10 nucleotide positions. The oral apparatus of tetrahymenids is morphologically plastic (right inset). Most species have maintained the plesiomorphic condition, while some have completely lost the oral apparatus (Clausilocola) or have evolved a highly complex, peritrich-like oral ciliature that was displaced posteriorly (hysterocinetids). Diagrams of Tetrahymena and Clausilocola in the right inset are from Zhang and Vďačný (2022, 2023).
Figure 10 in The search finds an end: the morphologically chimeric hysterocinetids belong to the subclass Hymenostomatia (Ciliophora: Oligohymenophorea)
Figure 10. Phylogenetic tree based on the nuclear 18S and mitochondrial 16S rRNA gene sequences, showing the phylogenetic position of hysterocinetids within the class Oligohymenophorea. Note that hysterocinetids are nested within the 'paravorax' clade of the genus Tetrahymena. The 'paravorax' clade, at the present state of knowledge, comprises both free-living (e.g. T. paravorax), as well as endosymbiotic ciliates associated with freshwater planarians (T. nigricans), bivalves (T. glochidiophila and T. unionis), and lumbricid earthworms (Hysterocineta bellerophon and Protoptychostomum simplex). Posterior probabilities for Bayesian inference conducted in MrBayes and bootstrap values for maximum likelihood conducted in IQ-Tree were mapped onto the 50%-majority rule Bayesian consensus tree. Fully statistically supported nodes are marked with red solid circles. The scale bar denotes one substitution per ten nucleotide positions.
Figure 7 in The search finds an end: the morphologically chimeric hysterocinetids belong to the subclass Hymenostomatia (Ciliophora: Oligohymenophorea)
Figure 7. Phylogenetic tree based on the 18S rRNA gene, showing the phylogenetic position of hysterocinetids within the class Oligohymenophorea. The class Colpodea was used as an outgroup, following Zhang and Vďačný (2022). Although hysterocinetids are morphologically highly dissimilar from Tetrahymena, they are nested within its 'paravorax' clade with strong statistical support (100% ML bootstrap, 1.00 posterior probability). The 'paravorax' clade thus became paraphyletic and its name-bearing species T. paravorax is depicted as a sister-taxon of both hysterocinetids with full support. Posterior probabilities for Bayesian inference conducted in MrBayes and bootstrap values for maximum likelihood conducted in IQ-Tree were mapped onto the 50%-majority rule Bayesian consensus tree. Fully statistically supported nodes are marked with red solid circles. The scale bar denotes eight substitutions per 100 nucleotide positions.
Figure 14 in The search finds an end: the morphologically chimeric hysterocinetids belong to the subclass Hymenostomatia (Ciliophora: Oligohymenophorea)
Figure 14. Secondary structure of the C, 3ʹM and 3ʹm domains of the 16S rRNA molecule of Hysterocineta bellerophon. Arrows mark molecular synapomorphies of the 'paravorax' clade. Similarly to the homologous nuclear 18S, molecular synapomorphies are distributed across the whole 16S rRNA molecule: 13 nucleotides in the C domain, seven in the 3'M domain, and two in the 3'm domain. Thus, 'paravorax' cladespecific nucleotide characters are situated in terminal loops of helices (three nucleotide positions), bulges and single-stranded regions (seven positions), as well as in the double-stranded regions (12 positions). As in 18S and 28S, many more mutations are involved in Watson‒Crick and wobble pairings than in non-canonical interactions in the double-stranded regions of the 16S rRNA molecule. The 16S secondary structure map of Escherichia coli (inset) is from hưp://apollo.chemistry.gatech.edu/RibosomeGallery.
Figure 4 in The search finds an end: the morphologically chimeric hysterocinetids belong to the subclass Hymenostomatia (Ciliophora: Oligohymenophorea)
Figure 4. Hysterocineta bellerophon in vivo. A, D, less-side overviews, showing the variability of body shape and size, as well as of the nuclear apparatus. B, detail of the anterior body region, showing the dense somatic ciliature and the unciliated sucker. C, detail of the vacuolized oral area. E, detail of the posterior body region, showing the oral ciliature and the vacuolized cytoplasm. CV, contractile vacuole; FV, food vacuoles; IF, infundibulum; MA, macronucleus; OC, oral cilia; S, sucker; SC, somatic cilia. Scale bars = 20 µm (E), 30 µm (B), 50 µm (A), 100 µm (D).
Figure 2 in The search finds an end: the morphologically chimeric hysterocinetids belong to the subclass Hymenostomatia (Ciliophora: Oligohymenophorea)
Figure 2. Protoptychostomum simplex in vivo (A, B) and asser protargol impregnation (C–I). A, detail of the anterior body region, showing densely arranged basal bodies of somatic kineties and the unciliated V-shaped sucker lined by regularly spaced, oblique rows of highly refractive granules. B, detail of the posterior body region, showing the oral ciliature and the vacuolized cytoplasm. C, D, less-side overviews, showing the variability of body shape and size, as well as of the nuclear apparatus. E, F, I, details of the infundibular and peristomial ciliature, as well as of the nuclear apparatus, which is composed of an ellipsoidal macronucleus and two globular micronuclei. Arrowhead in (E, F) marks the curved anterior end of membranelles M1 and M2. G, somatic kineties are composed of narrowly spaced monokinetids. Asterisks mark irregularities in the somatic ciliary paưern. H, surface view, showing cortical granules. CP, cytopharynx; F, fibre; G, granules; M1–2, membranelle 1 and 2; MA, macronucleus; MI, micronuclei; OA, oral apparatus; OC, oral cilia; PM, paroral membrane; S, sucker; SK, somatic kineties. Scale bars = 5 µm (I), 10 µm (F, G), 15 µm (A, B), 40 µm (E), 50 µm (C, D).
Figure 1 in The search finds an end: the morphologically chimeric hysterocinetids belong to the subclass Hymenostomatia (Ciliophora: Oligohymenophorea)
Figure 1. Protoptychostomum simplex in vivo (A) and asser protargol impregnation (B–I). A, less-side view of a representative specimen. B, F–I, less-side overviews, showing the variability of body shape and size as well as of the nuclear apparatus (shaded grey) and sucker (shaded yellow). Dashed circles in (I) represent contractile vacuoles. C, oral ciliary paưern. The paroral membrane and both membranelles extend along the whole posterior body end to plunge into the infundibulum where they form a helix-like paưern. Membranelle M1 is made up of two rows of basal bodies and is not segmented. Membranelle M2 runs beside M1, is composed of only a single row of basal bodies and is not segmented. Arrowhead in (C) marks the curved anterior end of membranelles M1 and M2. D, E, ciliary paưern of the less (D) and the right (E) side. CP, cytopharynx; CV, contractile vacuoles; M1–2, membranelle 1 and 2; MA, macronucleus; MI, micronuclei; PM, paroral membrane; S, sucker; SK, somatic kineties. Scale bars = 30 µm (I), 40 µm (A, B, D, E), 80 µm (H), 100 µm (F, G).
Figure 3 in The search finds an end: the morphologically chimeric hysterocinetids belong to the subclass Hymenostomatia (Ciliophora: Oligohymenophorea)
Figure 3. Hysterocineta bellerophon in vivo (A) and asser protargol impregnation (B–L). A, less-side view of a representative specimen. B, oral ciliary paưern. Membranelle M1 is made up of two rows of basal bodies and runs only on the peristome. Membranelle M2 is cut in two segments: (i) the distal segment M2 is built from two rows of basal bodies and runs beside M1, (ii) the proximal segment M2' is composed of three rows of basal bodies and extends beside M3. Membranelle M3 consists of two rows of basal bodies and starts at the infundibular entrance. C–F, I–L, less-side overviews, showing the variability of body shape and size as well as of the nuclear apparatus (shaded grey) and sucker (shaded yellow). Dashed circles represent contractile vacuole. G, H, the ciliary paưern of the less (G) and the right (H) side. Arrows mark a posterior secant system each on the ventral and the dorsal margin of the right body side. Abbreviations: CV, contractile vacuole; IF, infundibulum; M1–3, membranelles 1–3; MA, macronucleus; PM, paroral membrane; S, sucker; SK, somatic kineties. Scale bars = 50 µm (A, G, H), 100 µm (C, D, E, F, I, J, K, L).
Figure 6. Hysterocineta bellerophon asser protargol impregnation. A in The search finds an end: the morphologically chimeric hysterocinetids belong to the subclass Hymenostomatia (Ciliophora: Oligohymenophorea)
Figure 6. Hysterocineta bellerophon asser protargol impregnation. A, detail of the anterior body region of the specimen shown in Figure 5A, to document the organization of somatic kineties around the sucker. Skeletal fibres emerge from the anterior end of the less ciliary rows and run almost in parallel with the main body axis. B, detail of the oral ciliary paưern near the entrance to the infundibulum of the specimen shown in Figure 5C. C, D, membranelle M1 and M2 are made up of two rows of basal bodies each, run along the peristome, and terminate near the entrance to the infundibulum. Note that the posterior row of M2 is not recognizable in (C), as it is out of focus. E, F, paroral membrane extends on the opposite side of the infundibulum as M2ʹ and M3, describing 1.5 turns of a spiral. Arrowhead in (F) denotes the anterior end of M2ʹ and M3. G, somatic kineties are composed of narrowly spaced monokinetids. Asterisk marks an irregularity in the somatic ciliary paưern. H, the macronucleus is irregularly curved cylindroidal and contains innumerable small nucleoli. The small anterior blob is a preparation artefact caused by leaked macronuclear material as evidenced by the presence of nucleoli. F, skeletal fibres; M1–3, membranelles 1–3; MA, macronucleus; PM, paroral membrane; S, sucker; SK, somatic kineties. Scale bars = 3 µm (C, D), 10 µm (F, G), 15 µm (B), 20 µm (E, H), 30 µm (A).
Figure 9 in The search finds an end: the morphologically chimeric hysterocinetids belong to the subclass Hymenostomatia (Ciliophora: Oligohymenophorea)
Figure 9. Phylogenetic tree based on the nuclear 18S and 28S rRNA gene sequences, showing the phylogenetic position of hysterocinetids within the class Oligohymenophorea. The subclass Peniculia was used as an outgroup, following Gao et al. (2016). Posterior probabilities for Bayesian inference conducted in MrBayes and bootstrap values for maximum likelihood conducted in IQ-Tree were mapped onto the 50%-majority rule Bayesian consensus tree. Fully statistically supported nodes are marked with red solid circles. The scale bar denotes one substitution per ten nucleotide positions.
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