Rivularia auriculata, (VON MARTENS, 1875)

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, pp. 1-23 : 3-5

RIVULARIA AURICULATA (VON MARTENS, 1875)

Paludina (Melantho) auriculata von Martens, 1875a: 2 ; 1875b: 186

Organ system Specimens examined

Gross morphology All specimens (116176-1M, 116176-2M, 116176-3F, 116176-4M, 116176-5M, 116176-6F, 116176-7F, 116176-8M, 116176- 9M, 116176-10M, 116176-11M, 192196-1F, 192196-2M, 192196-3F, 192196-4F [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.

Rivularia auricularis – Heude, 1890: 178, pl. 40, figs 16 & 16a.

Rivularia auriculata – Kobelt, 1909: 178, pl. 35, figs 1–7; 12–16.

Type material

Lectotype ZMB 31048c, here designated ( Fig. 2A); 14 paralectotypes ZMB 31048a, b ( Fig. 2D).

Type locality

Siangkiang [Xiang] River, Hunan, China.

Remarks von Martens (1875a) 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 31048a and b, which correspond to the two varieties into which von Martens (1875b: 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, 16a) and reproduced by Kobelt (1909: pl. 35, fig. 1)]. Furthermore, the measurements reported by von Martens for two specimens match individuals from lot ZMB 31048a and b. Therefore, the largest specimen reported by von Martens is designated as the lectotype (from variety a; Fig. 2A) and is numbered ZMB 31048c. ZMB 31048a and b contain 14 additional specimens which then become paralectotypes ( ZMB 31048a, b; Fig. 2D).

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).