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Biodiversity Journal, 2019, 10 (3): 221–236

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Biodiversity Journal, 2019, 10 (3): 221–236 Biodiversity Journal, 2019, 10 (3): 221–236 https://doi.org/10.31396/Biodiv.Jour.2019.10.3.221.236 About the supra-generic classification of the Marginelliform Gastropods: a morphological study Franck Boyer 4, impasse Pasquier, 30190 Garrigues Sainte Eulalie, France: e-mail: [email protected] ABSTRACT The supra-generic classification of the marginelliform gastropods is reevaluated on the ground of a critical review of the main general revision works dealing with the combined analysis of the conchological characters, of the external anatomy and of the radular morphology, and less systematically of the organization of the alimentary tract. The disparate nature of several groups is demonstrated, and it provides grounds for the description of three new supra-generic taxa: Marginellonidae fam. nov., Plesiocystiscinidae fam. nov. and Canalispirinae subfam. nov. KEY WORDS Marginellidae; Cystiscidae; Granulinidae; taxonomic classification; morphology; cladistics. Received 26.08.2019; accepted 14.09.2019; published online 18.09.2019 INTRODUCTION fined as being composed of the “largest species”, with a radular plate said to be “flat or curved with The integrative taxonomy of the marginelliform many cusps”, whereas the Cystiscinae were defined gastropods has found probably its first hearty as being composed of “very small species with demonstration in the precursory work of Coan characteristic white shells” and provided with “a (1965) who proposed a relatively innovative clas- small and arched radular plate with fewer cusps”, sification confronting the conchological data with when the Marginelloninae were recorded as “rela- some available elements concerning the external tively large species with radular plates bearing nu- anatomy of the animals and the types of the radulae. merous cusps”. Coan (1965) proposed to recognize three sub-fam- The family Marginellidae did save his unifying ilies within a single family Marginellidae J. Flem- status, whereas the family Cystiscidae went down- ing, 1828: Marginellinae J. Flerming, 1828 and graded at the rank of sub-family within the Margin- Cystiscinae Stimpson, 1865 implicitely derived re- ellidae. In his conclusion, Coan (1965) detailed the spectively from Marginellidae and from Cystisci- composition of these three sub-families in genera dae Stimpson, 1865, as well as Marginelloninae for the fauna of the Panamic Province (“West Amer- subfam. nov., explicitely proposed for sheltering the ican species”), and he annexed a list of all the avail- two monospecific genera Marginellona Martens, able genera, as well as their synonymy. 1904 and Afrivoluta Tomlin 1947. The anatomical study of Ponder (1970) about In this formative period of the integrative tax- four marginelliform species from Indo-Pacific onomy of the marginelliforms, the definition pro- Province and its exploitation in the frame of a duced for each of these three sub-families remained broader reflexion about the taxonomy of the eminently summary: the Marginellinae were de- neogastropods (Ponder, 1974), followed by a work 222 FRANCK BOYER of Fretter (1976) about the Panamic species Volva- of operculum, polished surface of the shell, vitreous rina taeniolata Mörch, 1860 all together ushered a suture), Harasewych & Kantor (1991) report the true comparative anatomy applied to the marginel- convergent anatomic features which seem to sup- liforms, lifted the documentary investigations to port “the original assignation of this taxon (A’s N: higher level of detail and thoroughness, and pro- M. gigas) to the family Marginellidae (presence of vided the framework of a radical reconstruction of a buccal caecum, uniserial radula with broad the organization of the group. Ponder (1974) did rachidian “comblike” plate, wide aperture of the conclude about an intermediary position of the Mar- digestive gland, wide rectal gland and palleal ginellidae (sensu lato) between the Volutidae and oviduct with separated ingestive gland and seminal the Volutomitridae, while pointing out at the same receptacle)”, before to underline some significant time a greater proximity with the latter, based on divergency traits, especially the non-specialized al- the similarity of the columellar plaits and of several imentary system of M. gigas, with a strong Gland shared anatomic characters (one single ingestive of Leiblein without terminal bulb as well as an im- gland, right diverticula of the renal organ approach- portant Valve of Leiblein, which presents “features ing or penetrating the renal-genital duct, lateral of primitive neogastropods so far not-recorded in radular teeth lacking in the Marginellidae versus the Marginellidae”. Furthermore, M. gigas was said faint or lacking in the Microvolutidae, as section of to be “deprived of both a glandular duct bypassing the Volutomitridae). the Valve of Leiblein and as well as a salivary Directly inspired by these seminal works, gland, which are present in the documented Mar- Coovert (1986), continued by Coovert & Coovert ginellidae, including in the single species of Cys- (1990), did enter into a methodic work of documen- tiscinae studied (A’s N : for this feature) hitherto”. tation about the marginelliforms from various as- According to Harasewych & Kantor (1991), these pects, which will play as pieces preparing their particularities suggest that the group Marginelloni- “Revision of the Supraspecific Classification of the nae may constitute “the most primitive of the Mar- Marginelliform Gastropods” published on 1995. ginellidae subfamilies”, that means more primitive Among the most prominent of these preparatory than the Cystiscinae, considered itself by Coovert monographies: an essay on the genus Cystiscus (1989) as a primitive group within the Marginelli- Stimpson, 1865 (Coovert, 1986), another on the dae from the fact of the inclusion of a triserial radu- genus Afrivoluta (Coovert, 1987a), a compilation late species. about the external anatomy of the marginelliforms Coovert & Coovert (1995) published their (Coovert, 1987b), another compilation devoting to supraspecific revision of the marginelliforms, split- the conchological characters (Coovert, 1988), a ting the group in two families Marginellidae and third compilation about the general radulae Cystiscidae, the latter being restored for the occa- (Coovert, 1989), followed by a focus on the radulae sion. The Marginellidae received two subfamilies: in the Volvarina/Prunum group (Coovert & Marginelloninae Coan, 1965 and Marginellinae Coovert, 1990), displaying a good number of un- Flerming, 1828, the Marginelloninae remaining published data. composed of their two monospecific genera Mar- Harasewych & Kantor (1991) did extend the ginellona and Afrivoluta, and the Marginellinae Ponder’s method, applying it to the elusive species being distributed in three Tribes (Austroginellini, tr. Marginellona gigas (Martens, 1904) from south- nov., Prunini tr. nov. and Marginellini J. Fleming, eastern Asia. The authors confirm the belonging of 1828) constitued of many genera. Four sub-families Marginellona (and by extension: of the Marginel- were recognized in the Cystiscidae: Plesiocystisci- loninae) to the family Marginellidae, however with nae subfam. nov., Cystiscinae, Persiculinae subfam. some consistant reserves. After to have remembered nov. and Granulininae subfam. nov. This organiza- the initial attribution of M. gigas and of Afrivoluta tion was based on the comparison of relatively pringlei Tomlin, 1947 to the family Volutidae (in broaden compiled data concerning the shell mor- particular for the head shape and for the morphol- phology, the external anatomy of the animals (prin- ogy of the columellar plaits), and the original fea- cipally the structural shape of the head) and the tures which led to their subsequent placement in the morphology of the radulae, data completed for family Marginellidae (radular morphology , lacking some species and some genera by the description of About the supra-generic classification of the Marginelliform Gastropods: a morphological study 223 the radular odontophore and/or of the alimentary considered to be phyletically closer to the Margin- canal. ellidae and Volutidae than to the Cystiscidae. The This organization in families and sub-families principal traits adopted by Boyer (2017a) as diag- will be adopted as soon by the malaco-taxonomists nosis of the new family are: first the external and for the most it will remain undisputed up to anatomy (animal head of the Type 2 in Coovert & 2008, with the revision of the genus Serrata Coovert) and the pattern of the alimentary canal Jousseaume, 1875 for New-Caledonia by Boyer studied by Perez-Dionis et al. (2001) (“organ of (2008), who considers the Tribes organization pro- Leiblein made of a small bag provided with a short posed within the Marginellidae as a mainly eclectic duct rejoining the digestive tract after to have construction and suggests the transfer of the genera passed through the oesophageal nervous ring”), Serrata and Hydroginella Laseron, 1957 from the these two characters being considered as similar to Tribe Austroginellini G.A. Coovert et H.K. the pattern found in the Marginellidae, versus the Coovert, 1995 to the Tribe Prunini G.A. Coovert et slit head pattern occurring in the sub-families Cys- H.K. Coovert, 1995. Some new genera have been tiscinae and Persiculinae (the Type 2 animal being proposed also after the general revision of Coovert however found also in the Plesiocystiscinae, but & Coovert (1995), but without changing of the ar- constituting a plesiomorphic character,
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