Notes on the systematics, morphology and biostratigraphy of holoplanktic , 271. Comments on a paper discussing (, ) systematics, recently (2019) published in Bollettino Malacologico

Arie W. Janssen Naturalis Biodiversity Center, P.O. Box 9517, nl-2300 ra Leiden, The Netherlands; [email protected]

Janssen, A.W., 2020. Notes on the systematics, resent real Cuvierina, ancestral to modern Cuvierinidae. morphology and biostratigraphy of holoplank- The earliest real Cuvierina species, however, developed tic Mollusca, 27. Comments on a paper discussing from an ancestral Ireneia root during the late Oligocene/ Pteropoda (Gastropoda, Heterobranchia) system- early Miocene (Janssen, 2005). Praecuvierinidae should be atics, recently (2019) published in Bollettino Mala- retained as an unsuccessful offshoot of (presumably) Cre- cologico. Basteria 84 (1-3): 65-75. Leiden. Published 11 July 2020. seidae; (5) The relationship of the genus Vaginella with the Cuvierinidae, as suggested by Rampal (2019 and earlier), is denied and Vaginella is retained in the fam- ily; (6) The revival of the classic genusHyalaea Lamarck, Comments and critical notes are necessary concerning a 1799, is rejected. De Blainville (1821) is not its author and paper on pteropod systematics, published by J. Rampal in ‘Hyalaea’ cuspidata Bosc, 1802, is not its type species. 2019 (Bollettino Malacologico 55-2: 145-186). In that paper The nameHyalaea Lamarck, 1799, is a junior the author makes a number of statements on systematics of Cavolinia Abildgaard, 1791, with monotype Cavolinia and that cannot be ignored or left undiscussed. natans Abildgaard, 1791 = Anomia tridentata Forsskål The following issues are treated in this paper: (1) Notwith- in Niebuhr, 1775; (7) Phylogenetic relationships given by standing earlier discussions, the author maintains an ear- Rampal (2019) are based on cladistical and molecular anal- lier published interpretation of Cuvierina species which in yses that frequently are unclear or even contradictory. The the present paper (again) is demonstrated to be erroneous. molecular work was based on just two genes (coi and 28S). The unnecessary introduction of Cuvierina major Rampal, Divergence times of the various groups obtained from 2019, as a replacement name for C. atlantica Bé, McClin- ‘molecular clock’ interpretations usually differ strongly tock & Currie, 1972, is rejected, and the name Cuvierina from the fossil record evidence, mostly giving much older atlantica once more is accepted as valid; (2) The repeated datings; (8) The introduction in Rampal (2019) of the taxa argument to demonstrate the validity of the name Cre­ Heliconoididae and Thieleidae is accepted, be it with some seis acicula Rang, 1828, over C. clava Rang, 1828, is incor- doubt. The erection of Diacriinae andTelodiacria is con- rect and superfluous:Creseis clava cannot be considered sidered useful. A new family Hyalocylidae fam. nov. is nomen oblitum, and the validity of the name C. acicula for introduced herein. the type species of Creseis Rang, 1828, was decided by the First Reviser, d’Orbigny (1836); (3) The genus Altaspiratella Key words: Altaspiratella, Heliconoides, Creseis, Hyalo- Korobkov, 1966, transferred to the Pseudothecosomata by cylidae, Cuvierina, Praecuvierina, Vaginella, Hyalaea, Corse et al. (2013), repeated by Rampal (2019) is considered Diacriinae, Cavoliniinae, molecular analyses, molecular to be a true Euthecosomatous genus of the Limacinoidea. clock. Species of Altaspiratella show a gradual despiralisation leading to the first recognised Creseidae; (4) The Eocene family Praecuvierinidae Janssen, 2005, is rejected by Ram- pal, who considers the two genera of that family to rep- 1 For nr 26 in this series see Basteria 83 (4-6): 102-108 (2019).

basteria 84 (1-3): 65 A.W. Janssen – Comments on Rampal (2019, Bollettino Malacologico)

INTRODUCTION mentioned but two highly respected malacologists of the Muséum national d’Histoire naturelle, Paris, among others, Dr Jeannine Rampal, a biologist connected to the Aix-Mar- are thanked for ‘advice’ or ‘critically reading’ of the manu- seille University (France), published numerous papers on script. This was very surprising as it could not be believed systematics, morphology, biogeography and evolution of that these malacologists would have accepted, or even agreed present-day planktic gastropods. She graduated with a with some of Rampal’s opinions and decisions. Both of them, voluminous thesis (485 pp) on Mediterranean present-day however, did not read or approve the manuscript that was pteropods [Rampal, 1975]. More recently Dr Rampal coop- published in the Bollettino Malacologico (Ph. Bouchet, pers. erated with colleagues of the same university (Dr Emma- comm., 3 February 2020). Such a strong but untrue claim of nuel Corse and his team) on molecular research. Rampal does not contribute to the author’s credentials. Sur- The author of this paper is a retired curator of Ceno- prisingly, the manuscript passed the editorial procedures of zoic Mollusca of the former ‘Rijksmuseum van Geologie Bollettino, and was accepted for publication. en Mineralogie’ (currently incorporated in Naturalis Bio- diversity Center, Leiden. The Netherlands), but he is still active as an associate of that institute, specialising since the RESULTS early 1980s on predominantly fossil pelagic molluscs and producing multiple papers on systematics, morphology and Cladistics and molecular work biostratigraphy of that group of organisms. In her study the author at many places applies cladistic as Disagreements between Dr Rampal and this author have well as molecular analyses to discuss interrelationships and existed for a long time and have led to frequent email cor- phylogeny of species and higher taxa. Cladistics include respondence, but hardly ever to consensus. A recent paper 54 variables, 19 of which are based on shell characteristics (Rampal, 2019) in the Bollettino Malacologico requires a and the other on soft part anatomy. The molecular work is public rebuttal, as it gives rise to e.g. nomenclatural and almost exclusively based on Corse et al. (2013) and includes systematic confusions in this important group of Mollusca. sequencing based on two of the three in this respect pop- It is necessary to deal with it, since Pteropoda are currently ular genes, coi and 28S, the third gene that is frequently used in world-wide studies as indicators of e.g. the status of applied for molecular work, 18S, is not discussed. Conclu- marine ecosystem integrity. sions on relationships and phylogeny as presented in the 2019 Rampal paper are often unclear or even contradictory as e.g. (: 158) “Cladistic: Styliola is the sister group to Hyalo­ SOME HISTORY cylis and to the Cavoliniidae, – 28S mol. data (0.83/): it is the sister group to Hyalaea cuspidata and to the Cavoliniidae, The paper recently published in Bollettino has a long – 28S gene data set: it is the sister group to Hyalaea cuspi­ preceding history. An early version (Rampal [2014]) was data and to Cuvierina and ”. This probably is a result of submitted to ZooSystema. The editor invited this author the restricted number of analysed genes. A phylogenomic to review the manuscript, but as he never does a review approach based on transcriptome sequencing of multiple anonymously, and supposed that Dr Rampal would not be genes to fully resolve the phylogeny of pteropods will yield very happy with him as reviewer, he replied to the editor far better and more reliable data (Peijnenburg et al., [2019] only to be prepared to do so if Dr Rampal would accept and in review). The ‘molecular clock’ is frequently used to him as reviewer. As expected, this was not the case, and estimate divergence times in geological history. Results therefore, the manuscript was reviewed by someone else. of this method often differ strongly from what is known This apparently resulted in a rejection or withdrawal of the from the fossil record factual information and usually give manuscript: it never appeared in ZooSystema. much older dates than the fossil evidence does. Therefore, A drastically changed version of that manuscript (Ram- estimated divergence times based on the molecular clock pal, 2017) appeared in BioRxiv, which is an online internet should be treated with caution. archive accepting preprints of manuscripts without peer reviewing. Papers in BioRxiv are invalid for nomenclatural Columellar and umbilical morphology of gastropod purposes and taxa described as new therein are unavaila- shells ble. The 2017 version, therefore, was not indexed for Mol- Rampal (2019, e.g.: 147) seems to misunderstand the mor- luscaBase/WoRMS. A version with practically the same title phology of a spiral gastropod shell, as is clear from the has now formally been published in Bollettino Malacolog- expression “spiral calcareous shells with an umbilicus (but ico, but the various new taxa cited with the year 2017 are only no columella)”. Generally speaking, a gastropod shell will validated in this paper and, therefore, should be dated 2019. consist of a number of whorls increasing in diameter from In the Rampal (2019) Acknowledgements no reviewers are the apex downwards in a spiral around a central axis. If

basteria 84 (1-3): 66 A.W. Janssen – Comments on Rampal (2019, Bollettino Malacologico) the spiral is tight the most central portions of the whorls who only wrote (1992: 14) about Altaspiratella bearnensis together will build a massive axis (or columella) and there (Curry, 1982): “... a twisted columella, an anterior sulcus, will be no umbilicus at the base of the shell. If the spiral and a broad outer lip”’, agreeing with the original descrip- is less tight, the inner parts of the whorls will not touch, tion of Curry (1982) and with actual specimens. The word resulting in an open axis (or columella) and there will be ‘rostrum’ was nowhere used in the text of Hodgkinson et a basal umbilicus (that can secondarily be covered again, al. (1992). Indeed, in Altaspiratella an ‘anterior rostrum’ is as e.g. in Neverita). In shells in which the inner walls of the not present, just a more or less strongly accentuated notch whorls have an even greater distance from the central axis on the lower part of the thickened apertural margin, accen- these parts will form a wide open columella that allows a tuating the sinus between that point of the margin and the view into the interior from the base of the shell upwards, as base of the last whorl, not at all connected with the twisted is the case in the various despiralising Altaspiratella limac- columella proper. This feature is most clearly developed in inid species and will appear as a twisted 3-dimensional spi- Altaspiratella multispira (Curry, 1982) and is also present ral unit and not form an umbilicus as usual. on the basal apertural margin of Camptoceratops priscus (Godwin-Austen, 1882) and C. americanus Garvie, 1992, Systematic position of the genus Altaspiratella the first Creseidae species developing from the despiralis- Korobkov, 1966 ing limacinids. A similar notch on the apertural lip is now The discussion on the systematic position of the genus also known from another limacinid, Heliconoides nikkiae Altaspiratella ( or Pseudothecosomata) Janssen, 2017, from the Eo-Oligocene of Tanzania (Cotton was started in Corse et al. (2013, table 3) and is repeated in et al., 2017). Though in some species of Peraclidae a twisted Rampal’s (2019: 149) paper, with argumentation to reject columella is present, this never is in a wide, three-dimen- Altaspiratella as belonging to the Limacinoidea and to sional torsion as in Altaspiratella. The Peraclidae columella demonstrate it as a Pseudothecosomata species related to is tight and forms no umbilicus, not even in species that the Peraclidae. Earlier authors, however, all considered are wider than high, and there is no possibility to observe Altaspiratella (or its synonym Plotophysops Curry, 1982) as the shell’s interior, as in Altaspiratella species, in which the a limacinid: Korobkov (1966); Curry (1982); Hodgkinson et columellar torsion shows successive stages of despiraliza- al. (1992); Cahuzac & Janssen (2011), Janssen (1990, 2003); tion. Furthermore, apertural reinforcements like thicken- Janssen & Peijnenburg (2013, 2017), Garvie et al. (2020). ing of the margin or a basal notch are absent in Peracli- The type species of Altaspiratella, Physa elongatoidea dae, in which, on the contrary, some of the species develop Aldrich, 1887, for a long time was only known by its holo- apertural spines, as well as a basal columellar rostrum, all type, an incomplete specimen without apertural features of which are absent features in the various Altaspiratella preserved. Recently, however, well-developed specimens species. of that species were recovered, showing similar features as There also is the enormous difference in age: Altaspi­ present in other Altaspiratella species: a three-dimension- ratella ranged during Ypresian and Lutetian, whereas the ally twisted columellar spiral; a thickened, reinforced aper- oldest recognised Peraclidae species is known from the late tural margin; and a notch at the base of the apertural mar- Oligocene (Janssen, 2012b: 447), an age difference of about gin (Garvie et al., 2020). 20 million years. Where were they during that, for ptero- Based on Corse et al.’s (2013) opinion, Rampal (2019: pods well-investigated interval? 149) provided the following reasoning to demonstrate the Finally, the successive stages of despiralization in non-limacinid character of the genus Altaspiratella: “One Altaspiratella, causing the presence of an open columel- particulary striking feature which distinguishes Altaspi­ lar spiral and leading to the first Creseidae Camptoceratops ratella from the Limacinidae is the absence of the umbilicus and Euchilotheca, can barely be denied and was basically and the presence of a twisted columella ending in an abap- already for a long time accepted by various authors among ical rostrum with a narrow columella membrane (Hodg- them also Rampal [1975] and Curry & Rampal (1979). kinson et al., 1992)” and “The columella itself is thickened Accepting a transfer of Altaspiratella to the Pseudothe- and shows a distinct torsion in such a way that looking into cosomata would inevitably lead to the very unlikely con- the shell’s interior is possible by a straight adapical view; clusion that Creseidae have developed from pseudotheco- the apertural margin is widened and internally reinforced somatous ancestors. with a rather solid ridge connected with the thickened col- umella” (Cahuzac & Janssen, 2010: 24)’. Distribution of Heliconoides mercinensis and Limacina Commenting on this statement we (Garvie et al., 2020) heatherae fail to find the expression “a twisted columella ending with Rampal (2019: 147) states: “Among the earliest known spi- an abapical rostrum” in Hodgkinson et al. (1992). Ram- ral shelled Euthecosomata the fossil Limacina mercinensis pal (2019) has invented the phrase, not Hodgkinson et al., Watelet & Lefèvre, 1885, emerged during the early Eocene

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(Ypresian) (56.5 my).” This species, however, was already is only a little different to the Atlantic entity (one base) but present during the Thanetian: “The species is known from coi confirms his difference (unpublished data). They give the latest Paleocene in Alabama, Tuscahoma Sand Forma- evidence to a clade with several species.”. tion, Bear Creek Marls) (Naturalis collection)” (Garvie et These conclusions confirm and extend Janssen’s (2004) al., 2020) and might even be considerably older. The one observations and may justify the introduction of a family and only Cretaceous pteropod known (Heliconoides sp., Heliconoididae Rampal, 2019. Since the year 2003 Janssen see Janssen & Goedert, 2016) of Campanian age (~ 80 Ma) considered all other limacinids with any form of apertural resembles H. mercinensis closely and is considered conge- reinforcement as belonging to the genus Heliconoides. This neric. is, not for the first time, criticised in Rampal (2019), but the Also, on p. 147 it says: “The fossilLimacina heatherae large variety of apertural structures in fossil limacinids Hodgkinson, 1992, is listed under Late Paleocene (Thane- does not (as yet) allow a more robust or useful subdivision, tian) (58.6 my)?” Garvie et al. (2020), however, question and it may be expected that ultimately a number of further this age, as the type material was collected from possibly genera will have to be introduced or validated next to Heli­ downhole contaminated samples of the Baton Rouge bore- conoides, of which Skaptotion Curry, 1965, most likely will hole (Louisiana, U.S.A.). The type specimen ofL. heatherae be allied. For the time being, and awaiting further subdi- closely resembles a European Oligocene (Rupelian) species, vision, the fossil Helicinoides species may conveniently be L. umbilicata (Bornemann, 1855). As Rupelian sediments considered Heliconoididae. are, indeed, penetrated in the same borehole the age of L. heatherae might be early Oligocene instead. Type species of Creseis Rang, 1828, and the Creseis clava-C. acicula discussion Number of species in recent and fossil Heliconoides In Janssen’s (2007a) paper on Philippinian Pliocene fossils, The type species of the genusHeliconoides d’Orbigny, 1836, it was stated that Creseis clava Rang, 1828, is the correct is a very common present-day pteropod species with a large name for the Creseis type species, rather than the com- geographical distribution. It is the only recent limacinid in monly applied name Creseis acicula Rang, 1828. I based my which some shell reinforcement occurs, restricted to the last decision on the page priority of C. clava (which admittedly adult whorl and the aperture. These structures were already is not a valid reason) and on the fact that Rang’s wording in mentioned by d’Orbigny and later authors (e.g. van der the descriptions was unclear, writing about C. acicula: “... Spoel, 1967: 51, figs 17-18), but were in more detail described c’est plutôt un variété” (“it is more likely to be a variety”[of by Janssen (2004), who distinguished two different mor- C. clava]). Dr Rampal, during long e-mail discussions, phologies in ‘Heliconoides inflata’ (the correct name should insisted on C. acicula as the correct name. These two points have been H. inflatus; iczn Art. 30.1.4.4). These morpholo- of view ultimately led to consulting the opinions of three gies were indicated as forms A and B, with reference to dif- (ex-) commissioners of iczn, who all three thought that C. ferent geographical distribution patterns of these morphs. clava could not be considered the valid name. Rampal also Present-day specimens from the Mediterranean all belong submitted a request to the Commission (iczn 2018), asking to form A, whereas Caribbean material represents form B. for a formal decision (Case 3758). The Commission, how- Intermediate, or maybe just difficultly recognisable speci- ever, returned the case as a decision of iczn was not neces- mens, occur in the central Atlantic Ocean. Later (Janssen, sary, stating that the question could be solved by means of 2012b) it was found that these two forms can already be the First Reviser rulings in the Code. That ruling, indeed, clearly distinguished in lower Miocene rocks of Malta, in was applied in Janssen’s (2018a) paper in which the valid- which they occur sympatrically. Such observations basi- ity of C. acicula over C. clava was accepted. His conclusion cally offer possibilities for a taxonomic separation, but so (2018a: 111) was: “It has to bе accepted that formally the far this has not been sufficiently investigated. There are соrrеct name for the Creseis tурe species is Creseis acicula also indications that at least one further form might be dis- and not С. clava, although I still fееl that this is against the tinguishable on shell characteristics in southern Atlantic, original meaning of Sander Rang. However, iczn ruling is present-day material (Burridge et al., 2017). decisive here”. On the basis of the various analyses, cladistics and molec- Rampal (2019: 157), however, twisted the full extent of ular, Rampal (2019: 151) concludes: “The presence of two the argumentation by quoting only “iczn ruling is deci- distinct lineages (1.00/100) in the Mediterranean and in the sive here”. Surprisingly, Rampal maintained her earlier rea- Caribbean Sea suggests the existence of different molecu- soning to declare C. clava as invalid, referring to iczn Case lar geographically isolated species. Our present molecular 3758 on which the Commission never took a decision. She analyses make out four several geographical species in the declared C. clava a nomen oblitum, referring to iczn Art. inflata group: Atlanto-Mediterranean, Indo-Pacific, South 23.9.1.2. But the argumentation is incorrect: to designate a Eastern Atlantic and North Indian species entity; this last taxon as nomen oblitum also Art. 23.9.1.1 has to be met, but

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Creseis clava was used after 1899 as a valid taxon of the spe- The genusStyliola Gray, 1847 cies group by Tesch (1913, 1948), Tokioka (1955), Chen & Bé Rampal (2019: 157) stated: “This genus may have been pres- (1964), van der Spoel (1967, 1976), and Janssen (2007), so it is ent since the Late Oligocene”. The ‘may be’ is not necessary according to the iczn not a nomen oblitum. The validity of here, as Styliola has repeatedly been recorded from Chat- C. acicula over C. clava, therefore, was decided by the First tian rocks, of the Mediterranean (Malta), France and the Reviser, which was d’Orbigny (1836: 123). North Sea Basin. Furthermore, Rampal (2019: 157) inexplicably cites the More problematic is the phylogenetical relationship of two names as: “Rang, 1828 described Cleodora (Creseis) Styliola with the other as expressed in several clava var. acicula”. If this would have been true indeed, the papers published during the last decade, including Corse whole long discussion about acicula/clava would have been et al. (2013), and repeated by Rampal (2019), demonstrating superfluous and would make C. clava, by the difference in that Styliola is not monophyletic with the other Creseidae, rank, automatically the valid name. But no, Rang (1828: 317, but without giving a firm solution. Similar remarks can be 318; the C. stands for Creseis, not Cleodora – see page 309) made concerning the genus Hyalocylis. Rampal concludes introduced both taxa at full specific rank:Creseis clava and for both that they should no longer be included in the Cre- Creseis acicula. seidae family, where they have been placed traditionally In Janssen (2012a), in a paper on East Mediterranean and suggests to consider both as incertae sedis. holo­planktic molluscs, he discussed the uncertainty about However, the general shell characteristics of especially the type species of the genus Creseis Rang, 1828 and who ‘Styliole’ recta (de Blainville, 1827) = Cleodora subula Quoy had designated it. Zilch (1959: 49) considered C. virgula & Gaimard, 1827, the type species of Styliola, are so close to Rang to be the type species, which was accepted by van Creseis that it cannot be separated from Creseidae, at least der Spoel (1967). Rampal (2002: 231), however, referred to as long as no better solution is offered by the biologists. C. acicula as the type and, indeed, it was found that Pelse- neer (1888: 45) had been the first to designateC. acicula as The genus Hyalocylis, the Hyalocylis protoconch, and the the type species of Creseis. Oddly enough, however, Rampal genus Praehyalocylis (2019: 155) without any comment again mentions C. virgula The genusHyalocylis Fol, 1875 includes a single accepted as the type. present-day species, its type species is Creseis striata Rang, 1828. Hyalocylis striata deviates more strongly from the Status of Boasia Dall, 1889 Creseidae architecture than Styliola by its extremely fragile, The monotypic taxon Boasia, introduced as a subgenus of slightly curved and annulated shell with an elliptical trans- Creseis, was based on the small present-day species Cleo­ verse section. Several related fossil taxa, such as Tentacu­ dora chierchiae Boas, 1886, originally described from the lites cretaceus Blanckenhorn, 1889, Praehyalocylis haitensis Pacific Ocean, off Panama. Its single species shows a few Collins, 1934 and Hyalocylis euphratensis Avnimelech, 1945, characteristics distinguishing it from other Creseidae, such were already synonymised (Janssen, 1999) with H. striata, as a regular annulation of the teleoconch, and a different but a Pliocene species from the Philippines, Hyalocylis mar­ shape of the protoconch. Even for specimens with lacking ginata Janssen (2007b: 71, pl. 3 figs 5-6; pl. 23 figs 9-11) differs or reduced annulations the species can easily be recognised basically by the possession of a clearly reinforced apertural if a protoconch is present. Such smooth, or almost smooth margin. morphs were described as ‘Creseis virgula constricta’ Chen The larval shell of H. striata remained unknown for a & Bé, 1964, from the northern Atlantic Ocean and are long time and several authors mentioned shedding of the known for example also from the Indian Ocean and Red protoconch and formation of a septum to close the apical Sea. Almogi-Labin, 1982 (a paper that Rampal (2019) twice opening. However, in the hundreds of specimens from sed- erroneously refers to as ‘Almoggi-Labin, 1882’, also in the iment samples studied by the present author an apical sep- references) stated that most Red Sea specimens belong to tum could not be observed. Bandel & Hemleben (1995: 231, the more or less smooth form, agreeing with my own obser- fig. 4e) explained this curious fact by describing an organic, vations (Janssen, 2007a: 152). uncalcified nature of the embryonic shell that will disap- Interestingly, this species is present in lower-middle Mio- pear rapidly after death of the . Still, calcification of cene sediments of southern Australia (Janssen, 1990: 28) this protoconch is also known, as Almogi-Labin (1982: 58) and it is also not rare in Pliocene deposits of Pangasinan mentioned many juvenile specimens from the Red Sea, and (Philippines) (Janssen, 2007b: 66), both, however, repre- from the Mediterranean Janssen (2012a) described speci- sented exclusively the constricta form. This might indicate mens preserved as internal moulds. It was concluded that that the fully annulated form is a quite recent development most probably calcification of the early shell parts depends in C. chierchiae. I do agree with a full generic status of Boa­ on local circumstances. sia, indeed. TheHyalocylis larval shell differs substantially from Cre-

basteria 84 (1-3): 69 A.W. Janssen – Comments on Rampal (2019, Bollettino Malacologico) seidae. There is an elliptical protoconch, about twice as distributed in the Atlantic Ocean and Caribbean, whereas high as wide, with a perfectly rounded tip. The transition C. columnella is restricted to the Indo-Pacific. to the teleoconch is a slight constriction, after which the differs from C. atlantica by a annulation of the teleoconch starts right away (Janssen, slightly larger shell height and the possession of micro-or- 2012a, figs 43h-l). namentation that is absent in C. atlantica. Rampal (2019) The illustration of aHyalocylis striata protoconch given still denies the larger size of C. columnella and insists that by Rampal (2019, fig. 12-O’) has nothing to do with that spe- C. atlantica also occurs in the Indo-Pacific. However, the cies. Its pointed apex and stronger constriction rather make interpretation of sizes was erroneous: when measuring the it Styliola. shell height of the Cuvierina columnella specimen illus- The group of taxa included in the genus Praehyalocylis trated by Rang (1827: 323, pl. 45 figs 1-8) as 4.9 mm, Ram- Korobkov in Korobkov & Makarova, 1962 (not Prehyalocy­ pal did not realise that the bar accompanying Rang’s draw- lis, Rampal, 2019: 170) is known since the late Eocene Pria­ ing represents actual shell height and not a 1 mm scale. The bonian (not ‘Lower’ Oligocene, Rampal, 2019: 170). It dif- specimen illustrated by Rang contrarily was 11 mm high, fers from Hyalocylis by its circular transverse section and not 4.9 mm. Ever since 2005 this discussion has contin- less curved shell. A larval shell illustrated in Korobkov & ued and therefore Janssen explained the whole situation in Makarova (1962, fig. 9) resembles the protoconch of pres- his 2018b paper, to which the reader is referred for further ent-day Hyalocylis closely. Therefore, it seems acceptable to details. consider Praehyalocylis ancestral to Hyalocylis. Consider- Rampal (2019), however, still maintains her points of view ing the various fossil occurrences the transition between and now also, although initially accepted, denies the valid- these two genera must have taken place during the early ity of Bé et al.’s name Cuvierina atlantica, saying: (2019: 161): to middle Miocene. As genetical differences betweenHya ­ “Bé et al., 1972 included the correct Cuvierina columnella locylis and the Creseidae are also considerable, as demon- atlantica in a calcareous shell study. They described C. c. strated by Jennings et al. (2010: 7, fig. 3), Corse et al. (2013: atlantica but they illustrated C. c. columnella: there is a mis- 9, figs 3-4), Burridge et al., (2017: 13, figs 2-3); Peijnenburg et identification”. This statement, however, makes no sense in al. ([2019] and in review) a family Hyalocylidae fam. nov. that specimens used by Bé et al. all were from the Atlantic, is here introduced to contain these two groups of species. where C. columnella does not occur. It just demonstrates that Rampal still has a wrong idea of the two Cuvierina spe- Cuvierinidae cies. These two species, however, are perfectly defined by Disagreements on Rampal’s and Janssen’s interpretations the C. columnella neotype and the C. atlantica lectotype. of the present-day Cuvierina species, C. columnella Rang, The decision to reject the name C. atlantica led Rampal 1827, and C. atlantica Bé et al., 1972 also have initiated long (2019: 161) to the unnecessary introduction of the name C. discussions by e-mail that, as usual, did not lead to any form major Rampal, 2019 as a replacement name for C. atlantica. of agreement. The latter taxon was originally described by The choice of the name major for the smaller of the two van der Spoel (1970) as Cuvierina columnella forma atlan­ species under discussion once more demonstrates incor- tica. Being introduced after 1964 at infrasubspecific rank rect interpretation. The long discussion of iczn ruling on p. that name was not available from its original publication. 161 makes no sense. Rampal (2019: 161) increases the effort Rampal (2002), therefore, introduced the taxon Cuvierina by stating: “Janssen (2005: 41, 45, 46, figs 8-11; 2019: 371) spoeli instead, but that name was introduced at full species described atlantica but illustrated columnella”. level, with a holo- and paratypes from the Indian Ocean, To avoid unjustified accusations of misidentifications in not from the Atlantic, where van der Spoel’s material came the Rampal (2019) paper it would have been useful if that from. Cuvierina spoeli, therefore, cannot be considered a author had taken a look at the relevant type specimens. replacement name or nomen novum, but contrarily is an By doing so, it would have been clear that the neotype of independent, available taxon of the species group. However, Cuvierina columnella also is a paratype of C. spoeli, and in Janssen’s (2005) paper on Cuvierinidae it was demon- that the lectotype of C. atlantica is also the holotype of the strated that the type lot of Cuvierina spoeli did not represent unavailable C. columnella forma atlantica. the same species as the forma atlantica van der Spoel, 1970, It was also pointed out (Janssen, 2018b: 372) that Cuvie­ but contrarily belongs to the Indian Ocean/Pacific species rina identifications in Corse et al. (2013, done by Rampal) C. columnella. In that same paper one of the paratype spec- were erroneous, as checked on their barcodes in GenBank imens of C. spoeli was designated neotype of C. columnella by Alice Burridge. Finally, Rampal, (2019: 160) lists several and furthermore it was demonstrated that forma atlantica localities in the Indo-Pacific, with coordinates, for three van der Spoel had been validated as a taxon of the species Cuvierina species, which includes ‘C. major’ (= C. atlan­ group as C. columnella atlantica by Bé et al., 1972. It was tica). All of these are places, where Cuvierina atlantica does furthermore ascertained that C. atlantica was exclusively not occur.

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Praecuvierina and Eocene ‘Cuvierinidae’ The genusHyalaea Hodgkinson et al. (1992: 31) considered several Paleogene One of the most curious and incomprehensible acts in the genera from northern America to belong to the Cuvierin- Rampal (2019: 154, 163) paper is the ‘revival’ of the long-for- inae: Bucanoides Hodgkinson, 1992, Loxobidens Hodgkin- gotten genus Hyalaea for which de Blainville, 1821, is con- son, 1992, and Tibiella Meyer, 1884. Two of their new species sidered the author and ‘Hyalaea cuspidata Bosc, 1802’ the were assigned to the genus Cuvierina, namely Cuvierina type species. gutta Hodgkinson, 1992, and C. lura Hodgkinson, 1992. It is unclear to which ‘de Blainville 1821, p. 9’ paper Ram- However, Janssen (2005) believes the first real Cuvierina pal is referring. In the two papers mentioned in the list of species develops from an Ireneia ancestor not earlier than references, one is a well-known paper in Dictionnaire des during the early Miocene. This first typicalCuvierina spe- Sciences Naturelles, vol. 22, p. 65 with the title ‘hyale, Hya­ cies, С. torpedo (Marshall, 1918), was described from the lœa. (Malacoz.)’ with in the first line of text: “M. de Lamarck early Miocene (‘Late Otaian’ = Aquitanian) of New Zea- est le premier qui ait établi, sous cette dénomination, un land. In this species the shedding of the larval shell occurs genre bien distinct”, which is a clear reference to Lamarck close to the protoconch, the septum, therefore, is small and (1799: 89), where the genus is introduced with the monotype the shell retains an elongate spindle-shape with а slender ‘Anomia tridеntаtа Forsk.’. The genusHyalaea therefore is a basal part, strongly resembling species of Ireneia, inclusive junior synonym of Cavolinia Abildgaard, 1791 (Abildgaard, of the presence of the longitudinal micro-ornament (Jans- 1791: 172), with monotype C. natans Abildgaard, 1791 = Ano­ sen, 2005: 29). For this reason and also for other reasons, mia tridentata Fоrskåll in Niebuhr, 1775. This makes clear like the strikingly small sizes and absence of micro-orna- that neither de Blainville, nor ‘Hyalaea cuspidata’ have any- mentation, it was not possible to retain the two northern thing to do with this genus. In the same paper de Blainville American Eocene species in Cuvierina and for them, in (1821: 93) stated about ‘Hyalaea cuspidata’: “ ... que l’on peut the same paper, two new genera were introduced: Praecu­ presque assurer positivement que ce n’est pas à cette famille vierina and Texacuvierina, respectively, and a new family, qu’elle appartient”. Praecuvierinidae, was erected. It was suggested also that A ‘re-establishing’ of a genus Hyalaea to include Clio cus­ the Praecuvierinidae were not ancestral to the later Cuvier- pidata makes no sense at all and Rampal’s act only leads inidae, but represented an earlier, unsuccessful offshoot to unnecessary confusion. The genus group nameBellardi ­ from (presumably) Creseidae. clio Janssen, 2004, was introduced to contain Clio cuspidata The other genera considered by Hodgkinson et al. to (type species!) and some related species. belong to the Cuvierininae are now transferred to the Cre- seidae. The occurrence of a basally truncated shell (as a Systematic position of the genus Vaginella Daudin, 1800 result of larval shell shedding) and the presence of a subse- As the genus Vaginella is exclusively known from the fos- quent basal septum, that, however also is known from Cre- sil record there are no possibilities for molecular work and seidae, such as Euchilotheca elegans Harris, 1894 (Janssen et phylogenetical interpretations therefore can only be based al., 2011), was why Hodgkinson et al. (1992) assigned these on shell characteristics. Just as in earlier papers Rampal taxa to the Cuvierininae. However, the transverse striation (2019: 162) insists on ‘close phenotypical affinities between as seen in species of Bucanoides or in Tibiella species, or the Vaginella and Cuvierina’: ‘In Vaginella several phenotyp- apertural structures as in Loxobidens and Tibiella species, ical features are strongly reminiscent of Cuvierina: a bot- are unknown in the Cuvierinidae but do occur in several tle-shaped teloconch with a central bulge, a dorso-ventrally Creseidae, thus supporting the transfer of Bucanoides, Lox­ depressed peristome (with an ellipsoïd to semi-circular obidens and Tibiella to the Creseidae (Garvie et al., 2020). shape) with a slightly concave ventral edge, a blunt apex Rampal (2019: 161) concluded: “The bottle shaped fossil ending in a septum, and a circular to sub-oval transverse C. lura that appeared early in the Middle Eocene may con- section depending on the species. stitute the original phenotype characteristic of the Indo- Although some of the argumentations seem to justify Pacific Ocean”. Rampal also stated (2019: 163) concerning Rampal’s point of view, a number of similar facts can be the two species of ‘Precuvierinidae’ (sic), that their assign- used to demonstrate a closer relationship between Vaginella ment in a separate family “is not very convincing for several and the Cavoliniidae: dorso-ventral compression, presence reasons, e.g.: these fossils appeared at the same time as the of lateral carinae, dorsal apertural margin higher than the other Cuvierininae fossils”. However, these Eocene ‘Cuvier- ventral one, shedding of larval shell is only present in two ininae fossils’, are all considered to belong to the Creseidae of the many Vaginella species, the lack of lateral slits occurs now. The origin of the genusCuvierina , as a linear descend- also in several Cavoliniidae (e.g. in Gamopleura), etc. But ant of Ireneia during the Late Oligocene/Early Miocene is the most important and quite convincing characteristic not contradicted by evidence provided by Rampal (2019, or demonstrating affinity with Cavoliniidae; the ‘ontogenetic earlier). criterion’ as Rampal indicated it after Cahuzac & Janssen

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(2010), but further left it out of consideration. This charac- They record the species from the Indian and tropical Pacific ter was extensively described (although at the time not fully Ocean, but de Blainville (1821: 81) mentioned as type local- understood) in Janssen (1985), demonstrating a drastic shell ity ‘l’Océan Atlantique, assez près des Barbades: lat. 23°, metamorphosis during the transition from larval to adult 36', longit. 27°, 22'. This will need further research to find specimen, very similar as what is known from Cavolinia out what exactly is Diacria quadridentata’, and thus, what and Diacria, but absent in Cuvierinidae. exactly is Telodiacria. In the case of the late Oligocene species Ireneia striato­ carinata Piehl, 2007, Cahuzac & Janssen (2010: 75) admitted Rampal’s 2019 ‘new classification’ that the presence of lateral carinae might suggest a com- Ever since the classification of Bouchet et al. (2017: 357), mon ancestor of Vaginella and the Cuvierinidae. This fea- based on the work of Klussmann-Kolb & Dinapoli (2006) ture occurs occasionally, e.g. it was also present in the late and Burridge et al. (2017), the classic Order Thecosomata, Oligocene/early Miocene genus Spoelia, also included in with two suborders (Euthecosomata and Pseudothecoso- the Cuvierinidae. But neither the earliest species of Vagi­ mata) was abandoned, and the Order Pteropoda, with three nella from the Rupelian, nor any other Vaginella species, suborders (Euthecosomata, Pseudothecosomata, Gymno- demonstrate similarities to Ireneia or any other cuvierinid. somata) was applied instead. Rampal (2019: 168) does not follow this and maintains Thecosomata, but without com- ment. Rampal (2019: 163, fig. 21) maintains the two subspecies of Rampal’s (2019) new families, Heliconoididae and Thie- Clio pyramidata, traditionally naming them ‘C. pyramidata leidae, are accepted here, although both taxa contain just pyramidata Linnaeus, 1767, and C. p. lanceolata (Lesueur, a single genus, each of them with a single present-day 1813)’. However, these two names are demonstrated to be described species. The reference in Rampal (2019: 153) ‘Thie- synonyms (Janssen et al., 2018). For the nominal subspe- leidae (Rampal, 1975)’ is of course incorrect as the taxon is cies a neotype was designated, whereas for the morph usu- introduced as new in the 2019 paper. ally indicated as C. pyramidata pyramidata, restricted to In the Cavolinioidea group the present author prefers to the North Atlantic, the name C. pyramidata angusta Boas, retain Cuvierinidae and Cliidae at full family rank and not 1886, was activated. as subfamilies of Cavoliniidae, as species in each of these Corse et al.’s (2013) estimate of the Cliidae age (37.8 Ma) groups offer so much mutual similarity in shell-morphol- seems correct. The earliest Clio species is known from the ogy, wheras each separate group displays massive differ- Bartonian (41.3-38 Ma), not from the ‘Lower’ Oligocene, as ences in comparison with Cavoliniidae s.str. A subdivision Rampal states. of Cavoliniidae into Cavoliniinae and Diacriinae is consid- ered useful. Cavoliniidae In Janssen (2012a), a paper on East Mediterranean holo­ planktic molluscs, comments were included on the taxon- ACKNOWLEDGEMENTS omy of Cavolinia species, as proposed in Rampal’s (2002) publication, as he could not agree with some of Rampal’s My good friend Jaap van der Voort (Ostercappeln, Ger- opinions and statements. Dr Rampal soon reacted with many) is thanked for linguistic help and useful comments. a new paper in the Bollettino (Rampal, 2014) in such an Three anonymous reviewers and the Basteria editor team offensive way that Janssen did not find it worth a reac- are thanked for their many remarks from which the man- tion, considering that all relevant facts had been given and uscript benefitted. thinking that every researcher interested in the topic could easily evaluate the correctness of these statements. 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