Bulletin of Zoological Nomenclature 69(3) September 2012

Transcription

1 Bulletin of Zoological Nomenclature 69(3) September Comment on Turbo bidens Linnaeus, 1758 (Gastropoda, CLAUSILIIDAE): request for setting aside the neotype (Case 3581: see BZN 69: 85 87) Dietrich Kadolsky Beneath the Surface Geoconsultants Limited 66 Heathhurst Road, Sanderstead, Surrey CR2 0BA, U.K. ( Welter-Schultes bases his application to set aside the neotype of Turbo bidens Linnaeus, 1758, as proposed by Kadolsky (2009), on a distorted presentation of the facts of the matter, which if taken on its own will make an informed decision of the Commission impossible. Statements fundamental to the case are presented as certainties when they are either non-unique interpretations or even contrary to the facts of the case, or they are omitted altogether. In Welter-Schultes paragraph (1) every sentence is factually incorrect: Linnaeus (1758, p.767) established Turbo bidens with a description and a bibliographical reference to a figure of a specimen collected in Italy near Florence (Gualtieri, 1742, Tab. 4, Fig. C) that did not match the description. Here, as well as throughout his application, he repeats the often published claim that Linnaeus (1758) meant Papillifera papillaris (Müller, 1774) when he established Turbo bidens. In order to reconcile the incompatibility of Linnaeus definition with the characters of Papillifera papillaris, authors claimed that Linnaeus descriptive term sutura subcrenata referred to the subsutural papillae of Papillifera papillaris, and that Linnaeus erroneously quoted Gualtieri s Figure C when he meant Figures D and/or E, which both illustrate Papillifera papillaris. In so doing, authors overlooked or ignored a) that Gualtieri s Fig. C does clearly show a subcrenulate suture (see Figs. 1, 3); and b) that Linnaeus did not mention the conspicuous colour pattern of P. papillaris shells which consists of a reddish brown subsutural band interrupted by white papillae (see Fig. 5). None of the authors who claimed that Turbo bidens was conspecific with Papillifera papillaris commented on why the subcrenulate suture depicted in Gualtieri s Fig. C should have been incorrectly described by Linnaeus with the words sutura subcrenata, or they even denied their existence, e.g. Schröter (1784). At the heart of this error lies the semantic failure of several authors to distinguish between the descriptive terms crenatus, a, -um (referring to the shape of the suture) and papilla (a rounded raised structure on the surface of shell); see also Giusti & Manganelli (2005, p.131). Consequently, there is no mismatch between diagnosis and cited figure. Further, Gualtieri did not provide a locality; Linnaeus gave merely southern Europe. Linnaeus (1767, p. 1240) repeated the information in the same form and added a reference to another figure which did match the description (Bonanno, 1684, Cl. 3, Fig. 41). The author s name is actually Buonanni. Linnaeus diagnosis does not cite the subsutural row of dots depicted by Buonanni, who may have lumped several species together: his text describes the brownish subsutural band and white papillae of P. papillaris, but his figure shows a shell with more numerous and smaller dots than P. papillaris has, which are, moreover, dark, and the brown subsutural band of P. papillaris is also lacking. Various Italian clausiliid species in the genera Siciliaria, Delima and Charpentieria have small white subsutural papillae. A fortiori, the

2 214 Bulletin of Zoological Nomenclature 69(3) September 2012 Figs Copy of Gualtieri s Fig. C (1742, pl. 3), cited by Linnaeus (1758) as an illustration of his Turbo bidens. The figure is rotated. 2. Cochlodina bidens (Linnaeus, 1758) (syn. C. incisa (Küster, 1876)). Neotype proposed by Kadolsky (2009). Shell height 16.4 mm. Florence, Giardino di Boboli (Museo di Storia Naturale degli Studi di Firenze, Sezione Zoologica La Specola, no ). Photos courtesy G. Manganelli (Siena). 3. Detail of Gualtieri s Fig. C, showing the subcrenulate suture in detail. 4. Detail of the specimen in Fig. 2, showing extremely fine vestiges of sutural crenulations. 5. Papillifera papillaris (Müller, 1774). Neotype proposed by Giusti & Manganelli (2005). Shell height 15 mm. Florence, Giardino di Boboli (Museo di Storia Naturale degli Studi di Firenze, Sezione Zoologica La Specola, no ). Photos courtesy G. Manganelli (Siena). reference is not part of the original publication of the nominal species Turbo bidens and hence irrelevant to its identity. The contradiction was discovered a few years later by Schröter (1784) who observed that Gualtieri (1742) had indeed given a figure of the species that matched the Linnean description but that was not Fig. C but Fig. E of the same plate. Schröter was the first to assume that Linnaeus (1758) had made a mistake and cited the incorrect figure. There is no contradiction, as already discussed. Schröter (1784, pp ) used for both crenulations and papillae the German term Einkerbungen

3 Bulletin of Zoological Nomenclature 69(3) September (literally, incisions ), and it seems that due to this inaccurate terminology he came to his erroneous conclusion. Furthermore, Schröter overlooked or ignored the fact that Gualtieri s Figure C does show a subcrenulate suture. In paragraph 3, Welter-Schultes cites Hanley s (1855) statements about a specimen of Papillifera papillaris being present in the Linnean collection as proof that Linnaeus did indeed mean Papillifera papillaris when describing Turbo bidens. Hanley actually writes: Of that genus [Clausilia] I found three or four species in the collection (some possibly added since), of which one alone agrees with the expression sutura subcrenata... Unfortunately Hanley did not identify the other two or three clausiliid species in the Linnean collection, and he, too, misapplied the Linnean term sutura subcrenata to a papillate suture. Considering the considerable mismanagement of Linnaeus shell collection described by Dance (1967) and acknowledged by Welter-Schultes himself, it remains uncertain whether Linnaeus syntypes of Turbo bidens were either a mixture of several clausiliid species including Papillifera papillaris, or whether P. papillaris was added later, either by Linnaeus himself or someone else. We can only interpret what has been written in the original publication of Turbo bidens, and this excludes P. papillaris from being considered as the species described and referenced by Linnaeus, Hanley s statements are also discussed by Kadolsky (2009, p ). In paragraph 4, Welter-Schultes states: This decision [i.e. Opinion 2176] implied that the name Turbo bidens should be used for Helix papillaris, the identity of which had nowhere been disputed. Article 80.5 of the Code states: An Opinion applies only to the particular case before the Commission and is to be rigidly construed; no conclusions other than those expressly specified are to be drawn from it. The ruling concerning Helix papillaris Müller reads: It is hereby ruled that the specific name papillaris Müller, 1774, as published in the binomen Helix papillaris, is not conserved. The name is not suppressed, nor is there any ruling affecting the identity of Turbo bidens Linnaeus, 1758 given. Considering that the Commission had been advised that (a) the nominal species Turbo bidens and Helix papillaris in their original concepts are not conspecific (BZN 62(3): 131; 63(2): ; 63(3): 200); (b) the neotype proposed by Falkner et al. (2002), apart from not agreeing with the original diagnosis and bibliographic reference of Linnaeus, 1758, was Gualtieri s illustration of a specimen (1742, Fig. E) of Papillifera papillaris without a location, which no longer exists. Thus, this neotype designation was invalid under the Code, Article 75.3 (BZN 62(3): 131; 63(1): 47); the ruling concerning Helix papillaris was understood by Kadolsky (2009) to mean that the Commission considered it unnecessary or unwarranted to suppress the name Turbo bidens Linnaeus, 1758, in order to protect the name Helix papillaris Müller, 1774, as proposed by Giusti & Manganelli, 2005, because either, any threat to the name Helix papillaris came only from misidentifications of the name Turbo bidens (such as that perpetuated by Falkner et al. (2002) and Welter-Schultes), and not from its real identity; or, if they were indeed synonymous, Turbo bidens should not be suppressed because it had been used in several modern publications as valid. This interpretation of the ruling obviously contradicts the statement in the abstract of Opinion 2176, which cites Helix papillaris as a subjective synonym of Turbo bidens. This statement can only be regarded as a misunderstanding or error, as nothing had

4 216 Bulletin of Zoological Nomenclature 69(3) September 2012 been submitted to the Commission to justify declaring Turbo bidens Linnaeus and Helix papillaris Müller subjective synonyms (see above). A fortiori, a pronouncement on subjective synonymy is a taxonomic judgment, which is outside the remit of the Code (Principles: items 1, 3 and 4). To ignore the Commission s statement about the subjective synonymy of Turbo bidens and Helix papillaris was therefore considered by Kadolsky (2009) as being in accordance with the Principles of the Code, even more so as this statement was not backed up by a neotype fixation for Turbo bidens in the ruling given in Opinion The neotype selection by Kadolsky (2009) was an attempt at a nomenclatural solution which honoured Opinion 2176 (rigidly construed) as well as the taxonomic meaning of the original definition of Turbo bidens Linnaeus, 1758 and to protect the well-known specific name Cochlodina laminata (Montagu, 1803) (see next paragraph). In his paragraph 6, Welter-Schultes states: Since the proposal of suppression of Turbo bidens was rejected and the identity of the taxon was nowhere disputed, there was no exceptional need for selecting a neotype (Article 75.3 of the Code). This statement is completely untrue. The varying identifications of Turbo bidens have been discussed, i.a. by Kadolsky (2009) which Welter-Schultes completely ignores. In summary, Turbo bidens Linnaeus, 1758 had been identified with the following clausiliid species: Alinda biplicata (Montagu, 1803) by Pulteney (1799); Clausilia bidentata (Strøm, 1765) by Montagu (1803) and Turton (1831); Cochlodina laminata (Montagu, 1803) by Müller (1774), Pennant (1777), Draparnaud (1805), Gray in Turton (1840), von Martens (1860), Pilsbry (1922), Forcart (1965) and many others, although mostly before 1848; Papillifera papillaris (Müller, 1774) by, e.g. Pfeiffer (1848) and the majority of subsequent authors until the paper of Forcart (1965). After that date, usages of the names papillifera and bidens for the same species coexist to the present day. Cochlodina sp., probably C. incisa (Küster, 1876) by Giusti & Manganelli (2005). Cochlodina incisa by Kadolsky (2009). Of the subsequent identification attempts of Turbo bidens that with Cochlodina laminata is significant. Müller was a contemporary of Linnaeus; he communicated with him and sent him specimens (Dance, 1967). It is conceivable that Müller provided Linnaeus with specimens of Cochlodina laminata, which he collected in Frederiksdal near Copenhagen, Denmark, or assured himself through inquiries from Linnaeus of the identity of Linnaeus Turbo bidens. Although in this author s opinion this is the most plausible scenario, it needs to be stressed that this is merely a possibility. For conclusions we are left to interpret only what was written by Linnaeus. H. Nordsieck (online, unpublished) points out that the shells of Cochlodina laminata (Montagu, 1803) and C. bidens (syn. C. incisa) are very similar, and cannot be differentiated by characters of the suture. While this is correct, it is unlikely that Gualtieri (1742) collected C. laminata in Italy, where it is restricted to montane regions (Manganelli, in litt. 2012). The name Cochlodina laminata has been placed on the Official List in Opinion 336 (Opinions and Declarations Rendered by the International Commission on Zoological Nomenclature 10: , March 1955); it is widely and uncontroversially used and would be protected by the neotype selection by Kadolsky (2009) for Cochlodina bidens.

5 Bulletin of Zoological Nomenclature 69(3) September Also in his paragraph 6, Welter-Schultes partly cites unpublished objections by H. Nordsieck (online, undated) against Kadolsky s neotype selection. Nordsieck declared Kadolsky s neotype designation invalid because the neotype specimen was alleged to disagree with Gualtieri s Fig. C cited by Linnaeus. Concerning the subcrenulate suture of Turbo bidens, Nordsieck passes over the fact that Gualtieri s Fig. C shows it, and argues intsead:... no author has ever described any species of Cochlodina in such terms [i.e. having a subcrenulate suture]. The argument is specious as a crenulate or subcrenulate suture results from the ribbing of the shell. If a shell is described as ribbed, there is no need to mention that the suture is crenulate as this is implied. Several species of Cochlodina are ribbed (contrary to Nordsieck s statements) and hence have a crenulate or subcrenulate suture. A fortiori, the writings of (unspecified) subsequent authors are irrelevant to the case. Linnaeus may have thought the character of the suture was peculiar because Gualtieri s figure shows faint crenulations but no ribs. The neotype of Turbo bidens (Kadolsky 2009: Figs. 3b, 4) shows vestiges of ribs which are strongest at the sutures and form very faint crenulations there. No doubt Gualtieri s crenulations are drawn inaccurately and exaggerated, but they do exist. Another reason given by Nordsieck to reject the neotype of Turbo bidens was that only Gualtieri s figure D, representing Papillifera papillaris, showed the two teeth in the aperture which Linnaeus mentioned in his diagnosis. This difference is obviously due to the inaccuracy of Gualtieri s figures: The apertural lamellae of Figures D and E, which both represent Papillifera papillaris, are drawn very differently in both cases and are unrealistic in both. This shows that the artist did not attempt an accurate depiction of the apertural lamellae of P. papillaris, and there is no reason to assume that the illustration of the Cochlodina species in Figure C is any more accurate. Also, no South European clausiliid species is known which shows exactly the lamellae configuration shown in Fig. C. In summary, Welter-Schultes aims to influence a decision of the Commission without disclosing the many facts which undermine his case seriously. Of course the Commission could still vote for Welter-Schultes request to make Turbo bidens and Helix papillaris objective synonyms, but then this should be done in the full knowledge of the facts. Such a decision requires the Plenary Powers to set aside the provisions of Article , and would replace the name P. papillaris (Müller) well established in current literature and without any dispute about its identity- with P. bidens which was certainly not Linnaeus species concept and about whose identity published opinion is divided up to this day. The only advantage of such action would be to prevent the name Cochlodina incisa (Küster, 1876) becoming a junior subjective synonym of Turbo bidens Linnaeus, The name Cochlodina incisa is not often cited in modern literature, so that Article 23.9 cannot be applied to it. The name P. papillaris has a far greater claim for protection than C. incisa. It is up to the Commission to decide whether the historical truth (i.e. the identity of Turbo bidens) should be arbitrarily disregarded to protect C. incisa, and the completely uncontroversial name Papillifera papillaris which is well established in modern literature is to be made a junior synonym of the name Turbo bidens, which throughout its long history up to the present day has been associated with conspicuously different species and erroneous interpretations.

6 218 Bulletin of Zoological Nomenclature 69(3) September 2012 An alternative solution is here submitted for consideration by the Commission, which would not require the use of plenary power: (1) to place the name Papillifera papillaris (Müller, 1774), as published in the binomen Helix papillaris and defined by a neotype (Giusti & Manganelli, 2005: 132, Fig. on p.133) on the Official List of Specific Names in Zoology; (2) to amend the entry on the Official List of Specific Names in Zoology of the name Turbo bidens Linnaeus, 1758 to read: bidens, Turbo, Linnaeus, 1758, Systema Naturae, Ed. 10, vol. 1, p. 767, as defined by a neotype proposed by Kadolsky, 2009 (pp , Figs. 3b, 4) (Gastropoda). Additional references Draparnaud, J.P.R Histoire naturelle des mollusques terrestres et fluviatiles de la France. 8, 165 pp., 13 pl. Paris. Falkner, G., Ripken, T.E.J. & Falkner, M Mollusques continentaux de France. Liste de référence annotée et bibliographie. Patrimoines naturels, 52: Paris. Martens, E. v Die Heliceen, nach natürlicher Verwandtschaft systematisch geordnet, Zweite Ausgabe, nach dem hinterlassenen Manuskript [von J. C. Albers] besorgt von Eduard von Martens. xviii, 360 pp. Wilhelm Engelmann, Leipzig. Montagu, G Testacea Britannica or natural history of British shells, marine, land, and fresh-water, including the most minute: systematically arranged and embellished with figures. xxxviii, 606 pp., 16 pls. J. White, London. Nordsieck, H., undated [post X. 2009]. Papillifera bidens (Linné 1758) (Clausiliidae, Alopiinae), a common, but little known species. Pennant, T British Zoology. 4. Crustacea. Mollusca. Testacea. 135 pp., index of plates, 93 pls. Benj. White, London. Pfeiffer, L Monographia Heliceorum viventium sistens descriptiones systematicas et criticas omnium huius familiae generum et specierum hodie cognitarum. 1(1): (1847) [before ]; (2): (1847); (3): (1848). Lipsiae. Pilsbry, H.A Types of Férussac s subgenera of Helix. Nautilus, 36(1): Pulteney, R Catalogues of the birds, shells and some of the more rare plants of Dorsetshire. From the new and enlarged edition of Mr. Hutchins s history of that county. 92 pp. J. Nichols, London. Turton, W A manual of the land and fresh-water shells of the British Islands, with figures of each of the kinds. A new edition, thoroughly revised and much enlarged, by John Edward Gray. 324 pp., 12 pls. Longman, Orme, Brown, Green & Longmans, London.