SPIRA 2007 Vol. 2 Núm. 3 Pàg. 191-196

Rebut el 8 d’octubre de 2007; Acceptat el 27 d’octubre de 2007

Reply to Quintana et al. (2007): Darderia bellverica Altaba, 2007 is the correct name for the Mallorcan fossil helicodontid

CRISTIAN R. ALTABA

Laboratory of Human Systematics, Universitat de les Illes Balears, 07071 Palma de Mallorca, Balearic Islands (Spain) E-mail: [email protected]

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For several decades, a very rare, take the occasion to criticize my work. enigmatic found in Quaternary Herewith I present a rebuttal of all their sediments on the island of Mallorca points. (Majorca) was attributed to Oestophora barbula (Rossmässler, 1838), a west- • The original description of Darderia Iberian species never recorded in the bellverica was published in February Balearics (Gasull, 1963; Cuerda, 1989). 2007. Although issue 26.2 of This identification was based on superficial Biodiversity and Conservation belongs similarity and incomplete preparation of the into the volume corresponding to 2006, few specimens available. A re-examination the actual publication date is clearly of the two shells belonging to the Gasull mentioned in both the print and online collection and now deposited at the Museu versions. de Ciències Naturals de la Ciutadella in • The claim by Quintana et al. (2007) that Barcelona (catalog number MZB 84–6550) their former paper was published in allowed the description of a new species 2006 is a factual misrepresentation. within a new endemic genus, Darderia The Bolletí of the SHNB has often been bellverica Altaba, 2007 (Figures 1 and 2). published with great delay relative to the publication date printed in recent Several months after this description issues; volume 49 is no exception. was published in Animal Biodiversity and Indeed, on the cover and frontispiece of Conservation (Altaba, 2006, published in this issue the date printed is December 2007), a paper has appeared in the Bolletí 2006 (on pages 11 and 13, the date is de la Societat d’Història Natural de les 2005). This date cannot correspond to Balears (Quintana et al., 2006, published in the actual publication date for the 2007), where the same taxon is described following reasons. First, there are as a new species, Oestophora cuerdai several papers stating that they were Quintana, Vicens et Pons, 2007, on the accepted on “29-des-06”; it is most basis of three additional specimens. Two of unlikely that any printer would produce these belong to the Cuerda collection, and distribute a printed volume in just currently located at the Societat d’Història two days. Second, this issue includes Natural de les Balears (SHNB) in Palma. (p. 213) an obituary for someone Now Quintana et al. (2007), the authors of deceased in late January 2007 (Moyà- this binomen, claim to have priority and Solà & Pons, 2006). This obituary is

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Figure 1. Holotype (MZB 84–6550A) of Darderia bellverica Altaba, 2007. Maximum diameter: 10.1 mm.

signed by one of the authors of of Quintana (2006) in volume 49 of the Oestophora cuerdai. Third, Darderia Bolletí. The recency of the last Bolletí is bellverica was included in the catalog of further confirmed by the fact that the Beckmann (2007: p. 91), together with summary alone is posted on the web an indication that another “Oestophora page of SHNB without the pdf files as sp. ind.” would be described in the previous volumes (SHNB, 2007). future by Quintana, Vicens and Pons. • It is worth noticing that one of the Beckmann’s book, to which Quintana authors of Oestophora cuerdai is himself contributed, was compiled in himself editor of the Bolletí. Thus, the just a few months and published in July claim by Quintana et al. (2007) that the 2007, in a hurry due to the author’s publication date of the description of terminal illness (as attested by the Oestophora cuerdai has to be biography on p. 255, dated “den 8. Juni considered 2006, simply because there 2007”). On p. 68 of the same book it is is no printed mention of a later date in stated that Quintana was working on it, is not only unfounded, but shows that another paper, which was finally journal’s irregular management of published in the same Bolletí of the publication dates—no matter how they SHNB vol. 49 (Quintana, 2006, invoke the Code (ICZN, 1999). published in 2007), which was just The preceding points lead to the started to be distributed after conclusion that, although there seems September. In summer, a pdf file of this to be no way of determining the exact paper circulated through the Internet, date of publication of Bolletí SHNB vol. with no page numbers and a heading 49, this issue was really published in belonging to a monograph of the SHNB late 2007, certainly much later than entitled “Quaternari i Geomorfologia. Animal Biodiversity and Conservation Homenatge a Joan Cuerda Barceló”. vol. 26.2. Oestophora cuerdai is thus a Such monograph was also cited by junior synonym of Darderia bellverica. Vicens et al. (2006), but it was never • The criticism of Quintana et al. (2007) published. Apparently a late decision on the description of Darderia bellverica was taken not to publish that is equivocal and misleading. The monograph, including at least the paper complex apertural dentition consisting

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Figure 2 Holotype (MZB 84–6550A) of Darderia bellverica Altaba, 2007. Maximum diameter: 10.1 mm. Reprinted from Altaba (2006).

of one angular and several palatal five infrapalatals) with the armature of denticles is already well described and the type species, D. bellverica (five figured (Altaba, 2006). The latter at infrapalatals, in contrast with other least are clearly visible in Quintana et species included in the genus). al. (2007: Fig. 1), who employ an • Likewise, the claim by Quintana et al. incorrect nomenclature to the apertural (2007) that there is a mistake in the armature of land snails (ibid.: p. 159): “it apertural angle measurement given in is only possible to observe a low Altaba (2006) is based on their failure angular tooth (or basal tooth) to correctly measure this angle. There accompanied by a very elongated is no discrepancy if the coiling axis is palatal tooth”. This is quite inexact for taken as the origin (as explicitly stated two reasons. First, an angular tooth is in the original description of Darderia not the same as a basal tooth – the bellverica). angular lies at the junction of the • Why Quintana et al. (2007) cast doubts parietal and palatal margins, whilst a on the allocation of Darderia within basal is by definition located at the Lindholmiolinae remains elusive. As distal end of the aperture. And second, already pointed in the original the palatals are not simply one description (Altaba, 2006), such elongated, but five: two basally fused placement depends critically on the into a nearly lamellar structure just interpretation of homologies in the under the peripheral keel, one apertural dentition and protoconch. The prominent at the lower angle, a small overall shape of Darderia bellverica is one between the previously mentioned, intriguingly reminiscent of and a more bulky, blunt, forward Hesse, 1931, a taxonomically isolated protruding thickening at the lower helicodontid genus comprising several angle. These palatals can be seen in species from the Eastern the figures given by Quintana et al. Mediterranean region. The only other (2006, 2007); why they fail to identify genus belonging to the Lindholmiolinae these denticles (and to recognize the Schileyko, 1978, is Gittenberger, angular) is unclear. Furthermore, 1968, a rare, monotypic endemic genus Quintana et al. (2007) are wrong in of eastern Iberia, whose aspect and trying to criticize the diagnosis of aperture are quite distinctive and Darderia in Altaba (2006), because they unusual. Nevertheless, the protoconch confuse the traits of the genus (one to miscrosculpture of Atenia (Martínez- SPIRA, Vol. 2 (3) 193 Associació Catalana de Malacologia (2007)

Ortí, 2006) is apparently identical to fundamental issue if our goal is to know that of Darderia, probably indicating the history of life through a coherent, phylogenetic proximity. Indeed, the low logical classification based on angular tooth of Darderia, a feature monophyletic groupings (Cela-Conde & absent from other helicodontids, could Altaba, 2002). The description of a new be homologous to the parietal lip of genus and species appears herewith Atenia. The latter feature is a justified under the premises of building homoplasy (Gittenberger, 1968) shared a that aims at reflecting with Trissexodon Pilsbry, 1895, which cladistic relationships and being belongs to a different subfamily, the phylogenetically informative.” Why Trissexodontinae (Nordsieck, 1987). Quintana et al. (2007) insist in pointing However, the parietal lip of Trissexodon at the absence of anatomical is continuous with, and can be characters remains obscure. At any interpreted as a prolongation of, its rate, their statement that the traits of flaring peristome. Thus, the mere Darderia are coincident with those of presence of a parietal tooth or lip may Oestophora is merely based on have a low phylogenetic value; yet, the superficial similarities. Darderia has existence of such an armature indeed a general habitus similar to extending from the angular area, in Oestophora Hesse, 1907, but differs in addition to the protoconch characters, having much more developed apertural may indicate a common ancestry of armature (including an angular tooth), Atenia and Darderia within the protoconch microsculpture lacking Lindholmiolinae. As a result of an distinctive spiral incised lines, and incorrect and incomplete interpretation showing hair pits in the teleoconch. of the denticles and protoconch, Various other extant helicodontid Quintana et al. (2007) claim that these genera also have a similar lenticular traits are widespread among the appearance, but have very different . Thus their criticism on aperture and/or spire. the biogeographical implications of • The inclusion of other fossil species homologies within the Lindholmiolinae from Western Mediterranean islands in is not applicable. Darderia is questioned by Quintana et • The description of new taxa based on al. (2007) on tenuous grounds. The shells alone is obviously unavoidable in incomplete nature of specimens from the case of fossil snails. The reasons certain localities is no reason to cast supporting the description of a new doubts on the discussion given in species and genus, together with the Altaba (2006). interpretation of homologous • The Plio-Pleistocene age of Menorcan characters, are explicitly given in the fossils mentioned by Altaba (2006) is original description (Altaba, 2006: p. criticized by Quintana et al. (2007), who 198): “A point needs to be made about state that they come from sites clearly the decision to create a new genus and belonging into the Pliocene. Whether species for the Mallorcan specimens. the term Plio-Pleistocene can be They cannot be placed within any employed to collectively refer to the known species, either Recent or fossil, Pliocene plus the Pleistocene is a without stretching the morphological semantic quiz beyond the scope of this variation of the recipient taxon far paper. At any rate, the age of the fossils beyond the limits known in extant from Menorca (Minorca) is surely not species. Likewise, the new species Miocene as originally reported by cannot be allocated into any of the Quintana (1995). currently recognized genus-level taxa, • According to Quintana et al. (2007), the without considerably blurring the holotype of Darderia bellverica is definition of the existing taxonomy. deemed incompletely figured and Describing a new genus is not theratological, and thus contrary to the convenient simply because its recommendations of the International hypothesized relationships are not with Code of Zoological Nomenclature. This Oestophora or any other superficially is not correct, because the only thing similar taxa. The dilemma between that was not figured in the original multiplying genera and moving species description of D. bellverica, as clearly among existing genera is not a purely stated in Altaba (2006), are tiny nomenclatural problem, but a remnants of encrusted hardened soil

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and a crevice in the lower surface. any will to share their findings. The Moreover, the claim that this specimen discrepancy with Quintana et al. (2007) is malformed contradicts any on this issue lies on their account and comparison of all known specimens of mine, but surely does not help the goals this species, and is just based on the of the SHNB. incorrect interpretation of apertural • The conservation state of the fossils dentitions by Quintana et al. (2007). In examined by Quintana et al. (2006, addition, it must be stressed that the 2007) is, according to them, better than provision in the Code against naming that of the holotype and paratype of theratological forms applies only when Darderia bellverica. This statement is the malformed nature of the holotype is subjective and open to critical explicitly recognized in the original examination. At any rate, the careful description—a situation that has process of cleaning described in Altaba nothing to do with Darderia bellverica. (2006) caused in no way any significant • Why Quintana et al. (2007) question the deterioration. Moreover, it is clear from type locality given in Altaba (2006) is the figures in Quintana et al. (2006, unclear. The original label in the Gasull 2007) that the holotype of Oestophora collection is clearly readable. cuerdai is damaged, even having an Furthermore, Quintana et al. (2007) open hole; both this specimen and the question the geologic setting given in accompanying paratype have that label and in Gasull (1963), simply encrustations throughout, remain only because the site has been destroyed partially cleaned and have badly eroded through housing developments. Why undersides. The claim by Quintana et Quintana et al. (2007) claim to know al. (2007) that Darderia bellverica best, even when admitting that they should have been based on the have been unable to find the fossil site, specimen(s) in their possession is remains unexplained. simply unfounded. • The new specimens mentioned by • In their final conclusions, Quintana et Quintana et al. (2007) are just one. The al. (2007: p. 162) boldly point at what comment in Altaba (2006) concerning constitute their own mistakes: “[it] must the absence of further findings be considered as an unfortunate obviously refers to my own experience, attempt where characters are utilized since I was not aware of their then and interpreted at the authors’ unpublished finding. In fact, Vicens et convenience”. I could not state it more al. (2006) only mention a work in press precisely. where an “Oestophora sp.” will be cited. Furthermore, this paper was published From all of the above, it can only be in Endins vol. 30, which was not concluded that the attempt by Quintana et distributed before March 2007 (as al. (2007) to sustain their supposed priority recorded in the library of the Universitat and to show the alleged superior quality of de les Illes Balears). their work is entirely flawed. The correct • With regard to the impediments to name for the Mallorcan fossil helicodontid is examine the Cuerda collection at the Darderia bellverica Altaba, 2007. This taxon SHNB, I asked several times to was carefully described and illustrated, and examine the specimens, or at least to Oestophora cuerdai Quintana, Vicens et know the catalogue numbers. As a Pons, 2007 is just a junior synonym of the member of the SHNB, I thought I had former (contra Quintana et al., 2007), based the right to use for scientific research a on an erroneous and incomplete collection recognized as a public interpretation of some characters. resource. Over the previous year, I was repeatedly told that the collection was REFERENCES still getting unpacked and catalogued. My intention was clearly and honestly ALTABA, C.R. (2006). A new land snail from stated in conversations with Quintana, the Quaternary of Mallorca (Balearic Vicens, Pons and other members of the Islands, Western Mediterranean): SNHB. Only after publication of the Darderia bellverica n. gen., n. sp. description of Darderia bellverica I was (, , told about the Cuerda specimens being Helicodontidae). Anim. Biodiv. Cons., studied by some foreign malacologists. 29: 195–200. Quintana and co-workers never showed BECKMANN, K.-H. (2007). Die Land- und

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Süsswassermolluken der Balearischen Systems der (Gastropoda: Inseln. Conchbooks, Hackenheim. ). Arch. Moll., 118: 9- CELA-CONDE, C. & ALTABA, C.R. (2002). 50. Multiplying genera versus moving QUINTANA, J. (1995). Fauna malacológica species: a new taxonomic proposal for asociada a Cheirogaster gymnesica the family Hominidae. S. Afr. J. Sci., 98: (Bate, 1914). Implicaciones 1-4. biogeográficas. Boll. Soc. Hist. Nat. CUERDA, J. (1989). Los tiempos Bal., 38: 95-119. cuaternarios en Baleares. 2nd ed. QUINTANA, J. (2006). Reconsideració Conselleria de Cultura, Educació i taxonòmica de Chondrula (Mastus) Esports del Govern Balear, Palma de fòssil de Mallorca i Menorca Mallorca. (Gastropoda: Pulmonata: Enidae). Boll. GASULL, L. (1963). Un nuevo molusco Soc. Hist. Nat. Bal., 49: 21-38. terrestre fósil para la fauna cuaternaria QUINTANA, J.; VICENS, D.; PONS, G.X. de Baleares. Oestophora (id.) barbula (2006). A new species of the genus Charp. Bol. Soc. Hist. Nat. Bal., 9: 81- Oestophora Hesse, 1907 (Gastropoda: 82. Pulmonata: ) from the GITTENBERGER, E. (1968). Zur Systematik Upper Pleistocene of Mallorca (Balearic der in die Gattung Trissexodon Pilsbry Islands, Western Mediterranean). Boll. (Helicidae, Helicodontinae) Soc. Hist. Nat. Bal., 49: 51-58. gerechneten Arten. Zool. Meded., 43: QUINTANA CARDONA, J.; PONS I BUADES, 165-172. G.X.; VICENS XAMENA, D. (2007). INTERNATIONAL COMMISSION ON ZOOLOGICAL Algunas anotaciones críticas sobre NOMENCLATURE (ICZN). (1999). Oestophora cuerdai Quintana, Vicens International Code of Zoological et Pons, 2006 (: Pulmonata: Nomenclature. Fourth Edition. The Helicodontidae). Spira, 2: 157-162. International Trust for Zoological SOCIETAT D’HISTÒRIA NATURAL DE LES Nomenclature, London. BALEARS (SHNB). (2007). Publicacions. MARTÍNEZ-ORTÍ, A. (2006). New anatomical http://shnb.org/publicacions.htm data on the Iberian endemic Atenia VICENS, D.; CRESPÍ, D.; GINARD, A.; GRÀCIA, quadrasi (Hidalgo, 1885) (Pulmonata, F.; PONS, G.X. (2006). Les coves litorals Helicodontidae). J.Conchol., 39: 55–60. situades a la franja costanera entre es MOYÀ-SOLÀ, S. & PONS, G.X. (2006). Rafael Mal Pas i el Cap Gros (Alcúdia, Adrover, paleontòleg (1911 - 2007) in Mallorca) (2a part). Troballa de restes memoriam. Boll. Soc. Hist. Nat. Bal., de vell marí (Monachus monachus) a la 49: 213-215. cova des Fonoll Marí. Endins, 30: 87- NORDSIECK, H. (1987). Revision des 100.

Nota de l’Editor.—El Comitè Editorial d’Spira considerà que l’article de Quintana et al. (2007) publicat en aquest número de la revista s’havia d’interpretar com una rèplica a l’article d’Altaba (2006), i per això el seu manuscrit es va acceptar condicionalment al fet de donar dret de contrarèplica al Dr. Altaba. El Comitè Editorial d’Spira ha mantingut en tot moment una posició neutral pel que fa als treballs de tots dos autors. Tot i que el Dr. Quintana forma part del Comitè Editorial d’Spira, pel que fa a la revisió del manuscrit del Dr. Altaba i del seu propi manuscrit, el primer fou exclòs del Comitè a tots els efectes.

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Editor’s note.—The Editorial Committee of Spira considered that the paper by Quintana et al. (2007) published in this issue of the journal must be interpreted as a reply to the paper by Altaba (2006), and hence their paper was accepted conditionally to giving right to counter-reply to Dr. Altaba. The Editorial Committee of Spira has always maintained a neutral position regarding the works of the two authors. Even though Dr. Quintana is part of the Editorial Committee of Spira, with regard to the revision of Dr. Altaba’s manuscript and his own manuscript, the former was excluded from the Committee to all effects.

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