A Revision of the Recent Species of Rissoina (Moerchiella), R. (Apataxia), R. (Ailinzebina) and R . (Pachyrissoina) (Gastropoda: Rissoidae)
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I I BULLETiND E L'INSTITUT ROYA L DES SCLE CES NATURELLES DE BELG IQUE, BIOLOG!E, 63:71-135, 1993 BULLETINVAN HET KONlNKLIJKBE LGISCH !NSTITUUT VOOR NATUU RWETENSCHAPPEN, BIOLOGIE, 63:71-135, 1993 A revision of the Recent species of Rissoina (Moerchiella), R. (Apataxia), R. (Ailinzebina) and R . (Pachyrissoina) (Gastropoda: Rissoidae) by Willy J.M. SLEURS Contents Resume Abstract .. 71 Les especes Recentes de Rissoina (Moerchiella), R. (Ailinzebina), R. Resume .. 7 1 (Apataxia) et R. (Paclzyrissoina) sont revisees. En total, 14 especes Introduction 7 1 sont reconnues. Deux nouvelles especes sont decrites : Rissoina Materials and methods 72 (Moerclz iella) kilbumi sp. nov. et R. (M.) okinawensis sp. nov. La Museum and collection abbreviations . 73 nomenclature ouverte est utilisee pour deux especes de R. (Ailinze Abbreviations used in figures 73 bina), qui sont probablement nouvelles pour Ia science. Pour chaque Taxonomic account ... .. .... 73 espece les caracteres de Ia coquille sont donnees, ainsi que, pour quel Rissoina d 'ORBIGNY, 1840 ... 73 ques especes, les caracteres de Ia radule, de l'opercule et de l'ana R. (Moerc/ziel/a) NEVILL, I 884 74 tomie. Pour chaque espece, Ia repartition geographique connue est R. (M.) antoni SCHWARTZ, 1860 74 don nee. R. (M.) artensis MONTROUZIER, 1872 79 Mots-cles : Gastropoda, R issoininae, taxonomie, anatomie, repartition R. (M.) dorbignyi A. ADAMS, 185 I 84 geographique. R. (M.) gigantea (DESHAYES, 1848) 92 R. (M.) kilburni sp. nov... .. 94 R. (M.) okinawensis sp. nov . 97 R. (M.) striata (Quov & GAIMARD, 1833) 99 Introduction R. (M.) strio/ata A. ADAMS, 1851 . 103 R. (Apataxia) L ASERON, 1956 . ... 105 The present paper forms part of a comprehensive study R. (A.) cerithiiformis TRYON, I 887 105 of the Rissoininae, which is currently in progress. R. (Ailinzebina) LADD, 1966 .. .. 112 R. (A .) abrardi (LADD, 1966) .. 11 3 Members of the subfamily Rissoininae differ from the R. (A.) bi/abiata BOETTGER, 1893 . 11 5 sister-group, the Rissoinae, in the spe1m duct of females R. (A.) e/egantissima d ' O RB IGNY, 1842 11 6 being separated from the lower oviduct gland for most of R. (A.) sp. A . .. 12 1 its length (PONDER, 1985; pers. observ.). Within the sub R. (A .) sp. B . ..... 122 R. (Pachyrissoina) BOETTGER, 1893 . 123 family the following genera are recognized as being R. (P.) pem·assa G. & H. N EVILL, 1874. 124 valid : Rissoina d'ORBIGNY, 1840, Zebina H. & A. R . (P.) walkeri SMITH, 1893 126 ADAMS, 1854 and Stosica BRUSINA, 1878. PONDER (ibid.) Discussion . 132 recognized the fo llowing subgenera within the genus Acknowledgements 133 References . 133 Rissoina s.l. : Rissoina s.s., Phosinella MbRCH, 1876, Rissolina GOULD, 1861, Pachyrissoina BOETTGER, 1893, Apataxia LASERON, 1956 and Buvignieria COSSMANN, 1921, the latter containing fossil species only. Additional data, especially with respect to their anato Abstract my, however, reveals that members of the subgenus Rissolina do not diffe r sufficiently from Rissoina s.s. The Recent species of Rissoina (Moerchiella), R. (Ailinzebina), R. species to consider it a distinct subgenus of Rissoina (Apataxia) and R. (Pachyrissoina) are revi ed. A total of fourteen s.s. However, as species of this group are assumed to be species are recognized. Two new species-group taxa are named : Rissoina (Moerchiella) kilburni sp. nov. and R. (M.) okinawensis closely related, they will be treatened in a forthcoming sp. nov. Two probably new species, that belong to R. (Ailinzebina), paper as a separate species-group of Rissoina s.s. On known from only a few specimens are described but not named. Shell the other hand, PONDER (ibid.) considered Moen·hiella features of all species are redescribed or described in detail and, where NEV ILL, 1885 and Ailinzebina LADD, 1966 to differ avalaible, data on the operculum, the radula and the internal anatomy is also provided. The known distribution of each species is reported. insufficiently from Rissoina s.s. species to warrant the Key Words : Gastropoda, Rissoininae, taxonomy, anatomy, geogra recognition of both Moerchiella and Ailinzebina as phical distribution. distinct subgenera. Additional data with respect to both I I 72 Willy J.M. SLEURS the radular structure - especially the morphology of the central tooth - and the internal anatomy, reveals that both groups differ in several aspects from Rissoina s.s. species and therefore they are considered here as dis tinct subgenera of Rissoina s.l. Species of the subgenus Pachyrissoina differ from Rissoina s.s. species in shell characters; however, as no data on the anatomy of species of the subgenus Pachy rissoina were available, the latter is only tentatively regarded here as a separate subgenus, pending the exa mination of the internal anatomy. The present paper attempts to review the Recent spe cies of the subgenera Moerchiella, Apataxia, Ailinze bina and Pachyrissoina of the genus Rissoina. Materials and methods Shells have been examined in and/or borrowed from institutions listed under Abbreviations. All shell fea tures were examined using a WILD M-5 stereomicro scope. Shell dimensions were measured by usi ng an ocular with a micrometer of an accuracy of 0.001 mm. As a rule only adult specimens with an intact proto conch were included, except in the case of immature type specimens. Shell measurements (Fig. 1) include the total shell length (L), the spire length (Ls) and the maximum dia meter (D). Furthermore, in sculptured species, the num ber of axial ribs on the last whorl (No.ax) and on the penultimate whorl (No.axp) were counted. Finally, the number of teleoconch whorls was determined (No. whorls), starting from the nucleus to the transition Fig. I. - Shell dimensions : D, maximum diameter; L, total protoconch/teleoconch, with an accuracy of 0.25 whorl. shell length; Ls, spire length. The protoconch is considered here as the entire shell formed prior to metamorphosis; the type of develop ment (planktotrophic, lecithotrophic or direct develop ment) was inferred from the morphology of the proto an extremely thin layer (30 nm) of gold. conch (JABLONSKI & LUTZ, 1980). The shell was removed by dissolving the latter in a The protoconch and the microsculpture of the teleo 10 % solution of HCI. conch were studied by using photographs from a Immediately · after dissolving the shell, the animal was PHILIPS scanning electron microscope (SEM). cleaned with distilled water and eventually transferred The operculum features are of little importance in the to a 75 % solution of methylalcohol. systematics of the Rissoininae; only the presence or The spirally coiled animal was uncoiled and pinned in absence of a peg on the inner side was used in the the dissection tank, with its inner side turned towards classification of the rissoinine genera. When opercula the observer. Observing the animal in this position, were available, the inner side was photographed. facilitates the examination of the relative position of Radulae have been examined by using SEM micro the internal organs (especially the genital apparatus). graphs. The radula was dissected from the buccal mass Before dissecting the animal, a drawing of the entire and transferred to a solution of KOH, where it re animal in this position was made with the help of a mained for a short time (usually 1 day) in order to camera Iucida. After studying the relative position of remove the surrounding tissues. Subsequently, the the internal organs, the pallial cavity was opened from radula was transferred into distilled water and even the middle of the anterior mantle edge to the posterior tually it was put into a small drop of distilled water on end of the mantle cavity. Further, the pallial gonoduct a cover glass. After the evaporation of the water, the of the female was removed from the animal and radula stuck to the cover glass, which was glued with observations were made of both the left and right side double tape on a SEM stub. The radula was coated with of the gonoduct. In male specimens, after determining '' A revision of the Recent species of Risso ina 73 the position of the penis in the mantle cavity, the penis SMF : Senckenberg Museum, Frankfurt am Main, was cut near its base, just behind the base of the right Germany cephalic tentacle, and several drawings in different USNM : National Museum of Natural History, Smith positions were made. sonian Institution, Washington D.C. After the study of the organs of the mantle cavity, the ZSI : Zoological Survey of India, Calcutta stomach was removed and the following parts were measured : the total length, the length of the style sac and the maximum diameter of the stomach (off the Abbreviations used in figures anterior chamber). In order to study the circumoesophageal ganglia, a cut a.c. anterior chamber of stomach was made from the anterior end of the snout to the b.c. bursa copulatrix anterior end of the neck. After removing the buccal c.m. columellar muscle mass and the oesophagus, the circumoesophageal gan d.g. digestive gland glia can be easily observed. The RPG ratio (length of e.m.c. end of the mantle cavity the supraoesophageal connective : length of the right f.p. fecal pellet pleural ganglion + length of the supraoesophageal con g.p. genital porus nective + length of supraoesophageal connective) h.g. hypobranchial gland (DAVIS e.a. 1976) and the LPG ratio (length of sub int. intestine oesophageal connective : length of left pleural ganglion k. kidney + length suboesophageal connective + length of l.c.ga. left cerebral ganglion suboesophageal connective) were determined, at least l.o.g. lower oviduct gland when the left pleural ganglion and suboesophageal l.pl.ga. left pleural ganglion ganglion were not abutting. l.p.t. left pallial tentacle All information about localities, mentioned m m.e. mantle edge "Additional material examined", was taken from the oes.