taxonID	type	description	language	source
039A6736915AFFAEFC79F9BFFA8892D1.taxon	materials_examined	Type species: Paludina auriculata von Martens, 1875, by subsequent designation (Crosse, 1891).	en	Bocxlaer, Bert Van, Strong, Ellen E., Richter, Romy, Stelbrink, Björn, Rintelen, Thomas Von (2018): Anatomical and genetic data reveal that Rivularia Heude, 1890 belongs to Viviparinae (Gastropoda: Viviparidae). Zoological Journal of the Linnean Society 182: 1-23
039A6736915AFFA8FC8DF8DBFBC491C5.taxon	description	Gross morphology All specimens (116176 - 1 M, 116176 - 2 M, 116176 - 3 F, 116176 - 4 M, 116176 - 5 M, 116176 - 6 F, 116176 - 7 F, 116176 - 8 M, 116176 - 9 M, 116176 - 10 M, 116176 - 11 M, 192196 - 1 F, 192196 - 2 M, 192196 - 3 F, 192196 - 4 F [BVB, ES, RR]) Operculum 116176 - 3 (RR); 116176 - 1 (BVB); 116176 - 6 (BVB); 116176 - 10 (BVB) Pallial organs (general) 116176 - 3 (RR); 116176 - 4 (RR); 116176 - 7 (BVB, RR); 116176 - 10 (BVB) Respiratory system 116176 - 3 (gill, osphradium [RR]); 116176 - 4 (gill [BVB]); 116176 - 5 (gill, gill leaflet [ES]); 116176 - 7 (gill and osphradium [BVB]); 116176 - 8 (gill and osphradium [BVB]); 116176 - 10 (pallial circulatory system; gill leaflet [BVB]) Alimentary system 116176 - 1 (radula [BVB, RR]); 116176 - 2 (gastric chamber, radula [BVB, RR]; gross morphology visceral alimentary organs [BVB]); 116176 - 3 (buccal apparatus, exterior and interior, digestive gland, gastric chamber [RR]; style sac, intestine [ES]); 116176 - 4 (buccal apparatus, exterior and interior, gastric chamber [BVB, RR]; buccal muscles, salivary ducts, radula sac, rectum [BVB]); 116176 - 5 (gastric chamber, radula [BVB, RR]); 116176 - 7 (buccal apparatus, exterior and interior [ES, RR]; gastric chamber [RR]; salivary glands [ES]; food groove [BVB]); 116176 - 8 (general alimentary gross morphology, food groove, buccal apparatus, buccal glands, mid-oesophagus, oesophagus, salivary glands, rectum [BVB]); 116176 - 10 (digestive gland, gastric chamber [BVB]); 116176 - 11 (gastric chamber [ES]); 192196 - 1 (radula [BVB, RR]); 192196 - 2 (buccal apparatus [RR]; radula [BVB, RR]) Reno-pericardial system 116176 - 1 (pericardium, rectal blood sinus, kidney, pores, kidney-ureter connection, heart [BVB]); 116176 - 3 (pericardium [RR]); 116176 - 5 (kidney, heart [RR]); 116176 - 7 (pericardium [RR]; kidney [BVB, RR]; ureter, kidney-ureter connection [BVB]); 116176 - 8 (pericardium, heart, ureter, kidney [BVB]); 116176 - 10 (pericardium, heart, kidney, circulatory system of the reno-pericardial system, ureter [BVB]) Nervous system 116176 - 4 (nerve ring [BVB, RR]); 116176 - 7 (nerve ring [ES, RR]; supra- and sub-oesophageal ganglia, visceral ganglion [ES]); 116176 - 8 (nerve ring and its position compared to the buccal apparatus, buccal ganglia, pedal nervous system; supra- and sub-oesophageal ganglea, statocysts [BVB]) Male reproductive system 116176 - 1 (testis, visceral vas deferens [BVB]); 116176 - 2 (penis, prostate [RR]); 116176 - 4 (penis, prostate, pallial vas deferens, testis [BVB, RR]); 116176 - 5 (general male reproductive anatomy, penis, prostate, proximal pallial vas deferens, testis, visceral vas deferens [RR]); 116176 - 8 (all pallial reproductive organs [BVB]); 116176 - 10 (testis [BVB]); 192196 - 2 (proximal pallial vas deferens, testis, visceral vas deferens [BVB]) Female reproductive system 116176 - 3 (general female reproductive anatomy [RR, BVB]; juvenile [RR]); 116176 - 7 (brood pouch, capsule gland, albumen gland, seminal receptacle, renal oviduct [BVB, ES, RR]; visceral oviduct, ovary [BVB, ES]; juvenile [BVB]) 192196 - 1 (brood pouch, capsule gland, albumen gland, seminal receptacle, renal oviduct, visceral oviduct [BVB]) Most organ systems were subjected to comparative studies in males and females. Individuals are identified by extending the six digit ZMB museum numbers with unique specimen numbers. On the first row (gross morphology) ‘ M’ or ‘ F’ indicates the sex of each specimen. The investigating author (s) for each organ system are indicated by their initials. Abbreviations for PCR conditions: D = denaturation, A = amplification, E = elongation. Paludina auriculata (Melantho) – von Martens, 1870 – 1876: 155, pl. 135, figs 4 – 6.	en	Bocxlaer, Bert Van, Strong, Ellen E., Richter, Romy, Stelbrink, Björn, Rintelen, Thomas Von (2018): Anatomical and genetic data reveal that Rivularia Heude, 1890 belongs to Viviparinae (Gastropoda: Viviparidae). Zoological Journal of the Linnean Society 182: 1-23
039A6736915AFFA8FC8DF8DBFBC491C5.taxon	materials_examined	Type material Lectotype ZMB 31048 c, here designated (Fig. 2 A); 14 paralectotypes ZMB 31048 a, b (Fig. 2 D). Type locality Siangkiang [Xiang] River, Hunan, China. Remarks von Martens (1875 a) did not report where the type material had been deposited in the original description; however, ZMB 31048 was part of von Martens’ collection, was collected by von Richthofen, the collector of the type material and is from the reported type locality. Furthermore, it was split into ZMB 31048 a and b, which correspond to the two varieties into which von Martens (1875 b: 186) subdivided the species. These lots also contain specimens that seem to have been used for the first illustrations of the species [von Martens (1870 – 1876: pl. 135, figs 4 – 6); Heude (1890: pl. 40, figs 16, 16 a) and reproduced by Kobelt (1909: pl. 35, fig. 1)]. Furthermore, the measurements reported by von Martens for two specimens match individuals from lot ZMB 31048 a and b. Therefore, the largest specimen reported by von Martens is designated as the lectotype (from variety a; Fig. 2 A) and is numbered ZMB 31048 c. ZMB 31048 a and b contain 14 additional specimens which then become paralectotypes (ZMB 31048 a, b; Fig. 2 D). As mentioned von Martens subdivided R. auriculata into two varieties: variety a with obsolete spiral sculpture and variety b with distinct spiral sculpture. Under Art. 72.4.1 of the Code, reference to distinct varieties could exclude specimens from the type series of a new nominal taxon. As all the specimens known to von Martens at the time were attributed to one of the two varieties, with no nominotypical form, the type series of R. auriculata comprises all the material mentioned and, hence, is eligible for lectotype designation. The lectotype and paralectotype illustrated here span the morphological variation of the sample (see Shell morphology). Further variations have been described in the literature, as forms (Smith, 1900), or subspecies (Kobelt, 1909). A number of additional species have also been assigned to Rivularia (Heude, 1890; Kobelt, 1909). Because of its characteristic shell morphology, Rivularia is a relatively well-defined genus (Heude, 1890) that has been maintained as distinct by most authors (Heude, 1890; Yen, 1943), although assignments to Vivipara, Paludina and other taxa were commonplace in the early literature (Bachmann & Gredler, 1894; Smith, 1900).	en	Bocxlaer, Bert Van, Strong, Ellen E., Richter, Romy, Stelbrink, Björn, Rintelen, Thomas Von (2018): Anatomical and genetic data reveal that Rivularia Heude, 1890 belongs to Viviparinae (Gastropoda: Viviparidae). Zoological Journal of the Linnean Society 182: 1-23
039A6736915FFFA1FCE3F965FD4A954E.taxon	description	Radula: The radula is small (on average ~ 2.3 mm but up to ~ 2.9 mm in length in adults), slender and delicate, taenioglossate with on average ~ 55 (but up to ~ 65) rows of teeth. The rachidian is symmetric and possesses a blunt central cusp which can be broad (Fig. 7 A), or narrow (Fig. 7 B – D); it is bordered by three to four pointed cusps on each side that decrease outwardly in size; the flanking cusps are typically longer than the central cusp. The lateral teeth (Fig. 7 A – D) are similar in width but longer than the rachidian, and somewhat asymmetrical. They have a slightly larger, blunter central cusp than that of the rachidian, which is bordered by four to five cusps on either side that decrease outwardly in size, again with the flanking cusps longer than the central cusp. The inner marginal teeth (Fig. 7 A – D) are more asymmetrical than the lateral teeth, with long slender shafts (half the width of the lateral teeth and rachidian) and a cutting edge with one large, rounded inner cusp, bordered by one to two inner pointed cusps, which may be lacking, and two to three outer pointed cusps. The outer marginal teeth (Fig. 7 A – D) have shafts that are approximately half the width of the inner marginal teeth, and are even more asymmetric with one outer rounded cusp and one or two pointed inner cusps. The most central of these pointed cusps is equal in length to the rounded cusp. Oesophagus: The anterior oesophagus emerges from the dorso-posterior aspect of the buccal apparatus and forms a simple, narrow, somewhat dorsolaterally flattened tube (especially in males due to the bulging prostate) that bears continuations of the dorsal folds and dorsal food groove; oesophageal pouches are lacking (Fig. 6 A). In the mid-oesophagus, which starts just behind the nerve ring, the continuation of the dorsal folds and the dorsal food groove display the effects of torsion, which are abrupt rather than gradual. The mid-oesophagus is somewhat glandular and curves broadly beneath the food groove; its walls are irregularly longitudinally ridged and glandular, but it does not produce outpocketings or pouches; it deflects towards the right at the posterior part of the mantle cavity to continue posteriorly along the columella, passing below the reno-pericardial complex. The posterior oesophagus is finely longitudinally ridged and widens just before opening ventrally to the posterior gastric chamber. Gastric chamber: The gastric chamber (Fig. 6 B) lies at the left aspect, just behind the elongate pericardium and embedded in the lobes of the digestive gland. It occupies ~ 0.5 whorls, and has thin, non-muscular walls. A prominent T-shaped ridge compartmentalizes the gastric chamber floor into three regions. The longitudinal limb (= glandular pad) emerges from the right, posterior gastric chamber wall and forms a ridge along the floor to the right of the oesophageal opening, which opens ventrally at the left, roughly at the level of the posterior gastric shield. No digestive gland duct opens in the vicinity of the glandular pad. At the anterior end of the longitudinal limb, the transverse limb crosses the gastric chamber floor just in front of the oesophageal aperture and bears the gastric shield to the right. A prominent sorting area is developed in the left gastric chamber floor and roof, anterior to the oesophageal opening and extends anteriorly to the intestinal groove. Two curving longitudinal folds in the gastric chamber roof border the left side of the sorting area; the inner represents an extension of the minor typhlosole, with a prominently curved anterior tip, and the outer has its anterior tip near the level of the anterior end of the gastric shield. The inner fold often diminishes posteriorly, but both folds continue into the recess that receives the oesophagus. Two additional longitudinal folds border the sorting area to the right, and extend posteriorly from the vicinity of the proximal end of the major typhlosole, i. e. the ciliated or marginal fold, and the second right fold. These folds are joined by a third fold that emerges from the anterior end of the gastric shield. All three of these folds curve posteriorly to the right into a recess that receives the oesophagus, and that deepens at the left of the longitudinal limb. The folds curve around the posterior margin of this recess and continue anteriorly to end in the roof of the gastric chamber at the height of the gastric shield. Another prominent ridge starts at the centre of the recess, opposite to the oesophageal opening and continues anteriorly in the roof with the other folds. Just to the left of the gastric shield, two additional elongated ridges run along the right side of the longitudinal limb for most of its length; they connect anteriorly to the rightmost longitudinal folds bordering the sorting area. Within the gastric chamber roof, from the prominent outer longitudinal fold to the left side of the main sorting area, a series of flattened finger-like ridges are elaborated. The ridges begin as small undulations at the anterior tip of the outer longitudinal fold, gradually elongating and broadening, spanning the roof and extending onto the gastric chamber floor into a deep trough behind the gastric shield at the right side of the glandular pad. Anteriorly the transverse limb of the T-shaped ridge separates a deep, concave, cuticularised style sac pocket in the right aspect; this pocket is interrupted ventrally to the left by the extensions of the major and minor typhlosoles and the sorting area. The transition between the gastric chamber and the style sac is demarcated by a sulcus that is bordered by two transverse ridges of which the posterior is more prominent. The sulcus and these ridges are more strongly developed at right, and in the right ventral aspect of this sulcus, two digestive gland ducts open; at the left a shallow gastric pouch is present within the sulcus. Digestive gland: The digestive gland is a large beige organ located in the visceral hump, usually occupying the ~ 1.5 apical whorls, with its anterior ~ 0.5 whorl supporting the right aspect of the gastric chamber (Fig. 4) and the posterior part of the pericardium. The digestive gland is drained by tubules that join the digestive gland ducts, which run along the columellar aspect of the visceral whorls; however, in males the testes overlie the columellar aspect of the digestive gland. The ducts increase in size anteriorly, and open to the gastric chamber via two openings in the sulcus between the gastric chamber and the style sac. Style sac: The style sac is a voluminous undivided sac that connects broadly to the gastric chamber and narrows strongly distally. Its ventral and right walls are embedded in the digestive gland. It bears two ventral typhlosoles that traverse its length, bounding the intestinal groove. Anteriorly the typholosoles curve towards the left and subsequently to the right to continue into the intestine; below the oblique limb of the major typhlosole a small, non-glandular caecum is elaborated (Fig. 6 C). A crystalline style is absent. Intestine: The intestine emerges from the anterior end of the style sac following a slight constriction, and curves ~ 180 ° around the distal end of the style sac to continue posteriorly; it overlies the pericardium and the right side of the style sac (Fig. 8 A). The typhlosoles of the style sac continue laterally into the intestine for its entire length. The intestine continues posteriorly and its posterior limit coincides with the posterior wall of the pericardium and the anterior part of the gastric chamber; there the intestine turns back on itself to widen into the much larger, thin-walled rectum. Rectum: The rectum is thin-walled and semitransparent so that the contents are easily visible externally (Fig. 5 A, B, D, E). Posteriorly the rectum borders the right dorsal side of the pericardium, and continues anteriorly at the right side of the mantle roof in the pallial cavity. Its walls contain abundant transverse ridges, and a ventral, longitudinal groove with thickened epithelium. The rectum is surrounded by a layer of connective tissue that thickens ventrally and is penetrated by blood vessels, including a prominent vessel dorsally at the left side, which comprises the rectal sinus. At the mantle collar, the rectum becomes more muscular, and terminates in a free anal papilla. The rectum typically contains faeces that are compacted into small, oval pellets.	en	Bocxlaer, Bert Van, Strong, Ellen E., Richter, Romy, Stelbrink, Björn, Rintelen, Thomas Von (2018): Anatomical and genetic data reveal that Rivularia Heude, 1890 belongs to Viviparinae (Gastropoda: Viviparidae). Zoological Journal of the Linnean Society 182: 1-23
039A67369155FFA1FF7BFE5AFC0496FC.taxon	description	Heart and circulation: The pericardium (Fig. 8 A) forms a voluminous, elongate chamber bounding the rear of the mantle cavity in the central to left aspect, and is strongly compartmentalized at its right ventral side by the anterior digestive gland. Dorsally to the right, the pericardium is bordered by the rectum, but at the far right the pericardium forms the body wall, and anteriorly at the far right it is separated by a membrane from the ureter that continues anteriorly. Dorsally along the mid-line and at the left it is bordered by the intestine and the style sac, whereas the left anterior side is bordered by the kidney, the posterior gill and food groove (Fig. 8 A). The heart lies antero-ventrally at the far left within the pericardium. The large auricle is somewhat anterior and to the left of the smaller, more muscular ventricle, and receives blood (= haemolymph) from a blood sinus at the base of the kidney (= the efferent vein of the nephridial gland), which is confluent with the afferent and efferent branchial veins, and via the efferent branchial vein it is connected to the auricle. The efferent vein of the nephridial gland is at the right continuous with a blood sinus at the base of the membrane separating the ureter from the pericardium, which is connected directly to the rectal sinus. From the left, posterior aspect of the ventricle a short constriction separates the ventricle from the bulbous aorta. The bulbous aorta produces two posterior visceral branches and the anterior cephalic aorta which passes to the left of the oesophagus into the cephalic haemocoel. The left branch of the visceral aorta penetrates the viscera shortly behind the ventricle below the style sac, whereas the right branch continues posteriorly along the floor of the pericardium at the anterior end of the digestive gland and further alongside the posterior oesophagus.	en	Bocxlaer, Bert Van, Strong, Ellen E., Richter, Romy, Stelbrink, Björn, Rintelen, Thomas Von (2018): Anatomical and genetic data reveal that Rivularia Heude, 1890 belongs to Viviparinae (Gastropoda: Viviparidae). Zoological Journal of the Linnean Society 182: 1-23
039A67369155FFA2FCE3FCC9FE56954E.taxon	description	The pedal and cerebral ganglia are connected by comparatively long, slender, curving cerebro-pedal connectives. Otherwise, the constrictions between cerebral and pleural ganglia and between pleural and pedal ganglia are limited and these ganglia appear to be fused rather than interconnected with connectives. There is no pleural commissure, nor are there zygoses between the pleural and sub- or supra-oesophageal ganglia. The connectives between pleural and oesophageal ganglia are long. The oesophageal ganglia lie alongside but slightly separated from the mid-oesophagus, roughly at the same antero-posterior position, but the supra-oesophageal ganglion is larger and more elongate than the sub-oesophageal ganglion. Long connectives from the supra-oesophageal ganglia continue posteriorly towards the single, small visceral ganglion. This ganglion overlies the oesophagus at the posterior end of the cephalic haemocoel; it produces a large nerve dorsally, and a smaller nerve at the left side of the oesophagus.	en	Bocxlaer, Bert Van, Strong, Ellen E., Richter, Romy, Stelbrink, Björn, Rintelen, Thomas Von (2018): Anatomical and genetic data reveal that Rivularia Heude, 1890 belongs to Viviparinae (Gastropoda: Viviparidae). Zoological Journal of the Linnean Society 182: 1-23
039A67369156FFBCFF49FE5AFF1F91AD.taxon	description	Male reproductive system: The large, light orange-brown testis lies posteriorly in the viscera, and is elaborated into two lobes (Fig. 9). The posterior lobe lies ventrally and in the columellar aspect of the posterior part of the digestive gland, whereas the anterior lobe occupies the columellar aspect of the anterior part of the digestive gland, and is expanded ventrally and dorsally. Both lobes lie in close proximity but are physically separated. Testis and digestive gland are adjacent, not interdigitated; both organs can be differentiated based on colour (light orange-brown and beige, respectively) and texture (lobules of the testis are smaller than the acini of the digestive gland). Small ducts drain both lobes and coalesce into the visceral vas deferens which lies along the columellar aspect of the whorls. Much variation exists in the width of the visceral vas deferens between individuals, but it is roughly half the diameter of the oesophagus. Anteriorly the visceral vas deferens lies dorsal to the oesophagus, and at the anteriormost margin of the anterior testis it connects to the pallial vas deferens which is ~ 0.3 whorl in length and proximally enlarged to ~ 5 × the width of the distal visceral vas deferens and glandular. The proximal pallial vas deferens lies adjacent to the columellar muscle to the right, and the anterior pericardium and posterior mantle cavity to the left, in a position comparable to that of the pallial oviduct in females. It is white and elongate, broader posteriorly and gradually narrows anteriorly. Its walls are thin, with the central lumen constricted by thick, interdigitating transverse glandular lamellae. At its anterior extremity, which lies at ~ ½ of the length of the mantle cavity, the proximal pallial vas deferens turns ~ 180 ° towards the left into the distal pallial vas deferens. The distal pallial vas deferens is shorter (~ ½ of the length of the proximal pallial vas deferens) and only slightly wider than the visceral vas deferens. It crosses the mantle floor to the proximal end of the bulging prostate which lies embedded in the mantle floor underneath the food groove. The glandular prostate spans more than ¾ of the cephalic haemocoel and deflects at the neck to the right, narrowing further as it continues into the dorso-ventrally flattened right tentacle, which functions as a penis. The prostate is constricted by thin, transverse, glandular lamellae. At its tip, the penis forms a tentacular hook that curves into a subterminal cavity when not in use. In contrast to the greyish dorsal surface of the penis, the tentacular hook is white, with a terminal gonopore. Female reproductive system: The small, lobate ovary lies along the columellar aspect below the anterior end of the digestive gland and is drained by a narrow, simple visceral oviduct (Fig. 10). The visceral oviduct joins the narrower, proximal, descending limb of the renal oviduct at its posterior junction via a broad duct that connects proximally to the complexly branched seminal receptacle. The seminal receptacle is positioned along the left side of the posterior brood pouch below the rectum and is triangular in lateral view with the narrow pointed tip projecting posteriorly. The organ is wedgeshaped in cross section, with the broad, curved dorsal aspect embracing the rectum, and tapering ventrally. The renal oviduct forms a thick, muscular, ciliated U-shaped tube embedded in connective tissue ventral to the brood pouch. A gono-pericardial duct is lacking. The distal, ascending limb of the renal oviduct is slightly wider in diameter than the proximal limb and opens after a sharp U-turn into a broad, flattened albumen gland, which also lies ventral to the brood pouch, just anterior to the midpoint. The albumen gland connects to the capsule gland, which is a slightly narrower, glandular tube that continues posteriorly embedded in the ventral surface of the brood pouch. The capsule gland opens to the brood pouch posteriorly to the right, and has been observed to contain encapsulated eggs at the junction. The brood pouch contains a longitudinal sperm groove, which extends from the posterior end of the brood pouch to the gonopore. The brood pouch is undivided, with thin, translucent walls; it typically contains four to five large eggs in a single row. Anteriorly, the pallial oviduct narrows abruptly, before terminating near the mantle collar in the vagina. The vagina is a free hanging, muscular tube with a round terminal gonopore. PHYLOGENETICS Based on its area of occurrence and its attribution by previous authors, Rivularia would be expected to be a bellamyine, but our phylogenetic reconstruction including various bellamyines indicates that Rivularia is a viviparine (Fig. 11). This result was unambiguously retrieved for all used methods of phylogenetic inference (i. e. maximum parsimony, maximum likelihood and Bayesian inference). The taxon comes out as sister group to European Viviparus in our phylogenetic reconstruction (Fig. 11). Furthermore, both of the included viviparid subfamilies form monophyletic clades.	en	Bocxlaer, Bert Van, Strong, Ellen E., Richter, Romy, Stelbrink, Björn, Rintelen, Thomas Von (2018): Anatomical and genetic data reveal that Rivularia Heude, 1890 belongs to Viviparinae (Gastropoda: Viviparidae). Zoological Journal of the Linnean Society 182: 1-23
