◄ Carnets Geol. 15 (4) ►
Contents
[Introduction]
[The historical concept ...]
[Lectotype of Belemnites sulcatus ]
[Status of Belemnopsis Edwards in , 1849, and Belemnopsis , 1878]
[Summary]
[Systematic palaeontology] and ... [Bibliographic references]
The University of the West Indies, Mona, Kingston 7 (Jamaica)
Published online in final form (pdf) on February 28, 2015
[Editor: Bruno ]
A reinvestigation of the validity of the belemnite genus Belemnopsis is undertaken, together with a survey of the early history of the usage of the names Belemnites sulcatus, Belemnopsis and Belemnopsis . Belemnites sulcatus has been variously equated with either Belemnites apiciconus or with the group of belemnites including Belemnites Altdorfensis and B. Beaumontianus . (and not ) subsequently designated a lectotype which may be valid and, in case it is not, is validated here. The species concept for Belemnites sulcatus, as based on this lectotype, places it in the genus Holcobeloides . Belemnopsis has date priority over Belemnopsis , but must be interpreted as an "incorrect original spelling" and, therefore, does not enter into homonymy according to the ICZN; Belemnopsis is thus a valid genus. subsequently nominated Belemnites sulcatus, which was figured as Belemnopsis sulcata by , and therefore is a valid designation because this species is amongst the original species included in Belemnopsis by . One of 's figures of Belemnopsis sulcata agrees with Belemnites apiciconus , but does not agree with Belemnites sulcatus as defined by its lectotype; as such this is a case of misidentified type species. Belemnites apiciconus , the species involved in the misidentification, is therefore designated type species of Belemnopsis here and validated by citing the ICZN. The actions taken here maintain nomenclature at the genus, family and suborder level in respect to the names Belemnopsis and Belemnosis and serve to stabilize the complicated nomenclature issues related to these taxa.
Belemnites (Belemnitida); taxonomy; International Code of Zoological Nomenclature; Belemnopsis ; Belemnopsis ; Belemnosis ; Belemnites sulcatus ; Belemnites apiciconus .
S.F. (2015).- A reassessment of the validity and affinities of Belemnites sulcatus , 1826, Belemnopsis in , 1849, and Belemnopsis , 1878.- Carnets Géol., Madrid, vol. 15, nº 4, p. 31-39.
Une réévaluation de la validité et des affinités de Belemnites sulcatus , 1826, Belemnopsis in , 1849, et Belemnopsis , 1878.- Une réévaluation de la validité du genre de bélemnites Belemnopsis est réalisée conjointement à un survol rétrospectif des premières utilisations des noms Belemnites sulcatus, Belemnopsis et Belemnopsis . Belemnites sulcatus a été indifféremment assimilé soit à Belemnites apiciconus , soit au groupe de bélemnites constitué de Belemnites Altdorfensis et de B. Beaumontianus . Ultérieurement, (et non ) a sélectionné un lectotype qui pourrait être valide et qui, pour le cas où il ne le serait pas, est validé ici. Le concept d'espèce pour Belemnites sulcatus, tel que fondé sur ce lectotype, permet de l'attribuer au genre Holcobeloides . Belemnopsis bénéficie de l'antériorité (date de priorité) sur Belemnopsis , mais doit être interprété comme une "faute orthographique originelle" et, par conséquent, ne saurait être considéré comme un cas d'homonymie selon le CINZ ; Belemnopsis est donc un genre valide. Ce fut ensuite au tour de qui désigna Belemnites sulcatus, qui avait été figurée comme Belemnopsis sulcata par , et qui est donc une désignation valide parce que cette espèce fait partie de celles incluses à l'origine dans Belemnopsis par . L'une des figurations de Belemnopsis sulcata par correspond bien à Belemnites apiciconus , mais pas à Belemnites sulcatus tel que défini par son lectotype ; ainsi il s'agit d'un exemple d'espèce-type mal identifiée. Belemnites apiciconus , l'espèce impliquée dans l'erreur d'identification, est donc choisie ici comme espèce-type de Belemnopsis et validée en invoquant le CINZ. Les mesures prises ici permettent de préserver la nomenclature au niveau du genre, de la famille et du sous-ordre en ce qui concerne les noms Belemnopsis et Belemnosis et de résoudre des problèmes nomenclaturaux complexes directement liés à ces taxons.
Bélemnites (Belemnitida) ; taxonomie ; Code International de Nomenclature Zoologique ; Belemnopsis ; Belemnopsis ; Belemnosis ; Belemnites sulcatus ; Belemnites apiciconus .
Belemnites sulcatus was erected by (1826, p. 59) with the description "Guard subcylindrical, elongated, having a longitudinal sulcus, and terminating in an acute apex" and was recorded from the "Inferior Oolite" from "Dundry, near Oxford". Two specimens were figured, 's Pl. VIII, figs. 3-4 and Pl. VIII, fig. 5. Following an extensive discussion, (1999) concluded that (1870), as First Reviser of Belemnites sulcatus , fixed the species concept and restricted the name Belemnites sulcatus to 's Pl. VIII, fig. 5, a form that was regarded by (1865, p. 5; 1870) as originating from the Oxford Clay (Callovian) from near Oxford. This conclusion has had serious implications for belemnite taxonomy (, 1999). Belemnites sulcatus was designated as type species of the genus Belemnopsis , 1878, by H. in 1879, but (1999, p. 60) maintained that the real Belemnites sulcatus , 1826, as emended by , 1870, is not represented amongst the species figured by (1878) and, as such, would represent an invalid designation. Furthermore, (1999) points out that (1926) recognised that Belemnopsis , 1878, was preoccupied by Belemnopsis in , 1849, and that therefore 's genus name was invalid. This leads to further complication because the family name Belemnopseidae , 1922, emend , 1946, and the suborder name Belemnopseina , 1965, are derived from Belemnopsis, whereas the family name Belemnoseidae , 1865, is based on Belemnosis. If Belemnopsis in , 1849, is valid then this genus would be placed under the Family Belemnoseidae creating extensive nomenclature confusion. Further, Belemnites sulcatus as emended by would be placed today in the genus Holcobeloides , 1958 (, 2011), which belongs to the Cylindroteuthididae , 1919. To reduce such nomenclatural complexity, (1999) suggested that Pachybelemnopsis (1980) should be used for those forms previously attributed to Belemnopsis, with the Suborder Pachybelemnopseina (in et al., 1998) and Family Mesohibolitidae , 1983, replacing Belemnopseina and Belemnopseidae, respectively. However, this course of action does not remove the problem of what to do with the generic names Belemnopsis in , 1849, and Belemnopsis , 1878 (depending on which one is valid), and some belemnite workers (e.g., & , 2007, p. 6) are unhappy with discarding so well-entrenched a generic name as Belemnopsis , 1878.
In this paper I explore the nomenclature problems surrounding Belemnites sulcatus using the rules of the International Code for Zoological Nomenclature (, 1999) and come to different conclusions. I present these arguments in this paper.
The exact composition of 's 1826 type series for his species Belemnites sulcatus is now impossible to determine, but some observations can be made. (1826) figured two specimens of Belemnites sulcatus, and therefore 's 1826, Pl. VIII, fig. 5 and 's 1826, Pl. VIII, figs. 3-4 clearly comprise part of the type series. In giving localities for his belemnite species, indicated different localities separated by commas; so that for Belemnites abbreviatus (1826, p. 59), we have two localities which were written as "Weymouth, Dundry", and which now would represent different species: a species of Pachyteuthis from the Oxford Clay or Corallian of Weymouth, and a species of Brevibelus from the Inferior Oolite of Dundry – yet both are recorded as coming from the "Inferior Oolite". For (1826), the "Inferior Oolite" at that time would appear to have included what we would now consider as Inferior Oolite (Aalenian-Bajocian) as well as the Oxford Clay (Callovian-early Oxfordian). The specimen illustrated in 's 1826, Pl. VIII, fig. 5 as B. sulcatus was presumably found "near Oxford" in the Oxford Clay (, 1865, p. 5; 1869, p. 101; 1870, p. 114), and 's 1826, Pl. VIII, figs. 3-4 illustration of B. sulcatus was presumably collected from the Inferior Oolite of Dundry, Somerset (, 1843, 177; , 1849, p. 136; , 1865, p. 5). (1826, p. 59) also refers to a specimen from the Oxford Clay of St. Clements that was illustrated in 's {sic } (1677, Pl. III, fig. 6) History of Oxford, and this therefore also qualifies as a syntype because (1826, p. 59) specifically makes reference to it. (1826) may have had more specimens available, but no other specimens are mentioned, nor preserved in museum collections as far as is known.
The first question to ask is if any of 's syntypes are still in existence? Mr. J.S. was from Gdańsk and resided in Bristol where he was curator of the Museum of the Institute in Bristol (, 1865, p. 5). He wrote various papers on fossils and much, if not all, of his collection was deposited in the Bristol Museum (Bristol Mercury, Tuesday 2nd November 1830, p. 4). It is therefore likely that most of the belemnites illustrated by in 1826 were in the collection of the Bristol Museum. It is notably that (1869, p. 101) stated that 's 1826, fig. 5 seems to be modelled on specimens that then existed (note the past tense even in 1869) in the Bristol Museum (which he states were labelled "B. sulcatus, Inferior Oolite"), and presumably 's 1826, figs. 3-4 was also in this collection. Unfortunately, the geology department and geological collection in the Bristol Museum were destroyed by bombing during the Second World War (Western Daily Press, Friday 6th December 1940, p. 5) and these specimens no longer exist. The whereabouts of 's 1677, Pl. III, fig. 6 is also unknown.
(1827, p. 68-69) discussed 's 1826 syntypes of B. sulcatus in relation to the new species that he erected in his memoir (see , 1999, for 's various earlier papers on belemnites). (1827) introduced two new species, Belemnites Altdorfensis and B. apiciconus, and unambiguously referred 's 1826, Pl. VIII, fig. 5 to the former, and 's 1826, Pl. VIII, figs. 3-4 to the latter. However he stated that he did not use 's name because there were at least five or six different belemnite species that had ventral grooves. As such, (1827) did not revise Belemnites sulcatus , or make an appropriate designation of a type specimen.
John produced the first edition of his Geology of Yorkshire in 1829. In this work (p. 138) he recorded Belemnites sulcatus from the Oxford Clay and Kelloways Rock. (1829, p. 117) stated that he referred to figures in his own work or to works of others; for Belemnites sulcatus he lists 's 1826, Pl. VIII, fig. 5. So even in 1829, 's may have been restricting 's Belemnites sulcatus to the form from the Oxford Clay, but there is no unambiguous indication of that in his publication. By the second edition of his work, published in 1835, he no longer listed Belemnites sulcatus from Yorkshire, and later stated (, 1870, p. 117): "I doubt the occurrence of the species [Belemnites sulcatus] in Yorkshire, and regard the mention of it in the first edition of my work on the geology of that county (1829) as requireing confirmation". (1829) clearly does not designate a lectotype for Belemnites sulcatus ( article 74.5).
Several authors discussed the relationship between Belemnites sulcatus and B. apiciconus in the 1840s. (1843, p. 177), in his Catalogue of British Fossils, restricted B. sulcatus to 's 1826, Pl. VIII, fig. 3, and listed it as occurring in the Inferior Oolite of Dundry, Somerset (thus excluding the reference to 'near Oxford'). (1843) does not refer to 's work on the Jurassic of France (which was published in parts from 1842 to 1847, and which described Belemnites sulcatus in 1843). It appears that (preface dated July 1843) had not yet seen 's work on the Jurassic. (1843, p. 105) also restricted Belemnites sulcatus to 's 1826, Pl. VIII, figs. 3-4 (excluding fig. 5 from his synonymy list) and stated that this form was originally described by (1823, sic 1826) under the name Belemnites sulcatus and that (1827) changed the name to Belemnites apiciconus. (1843, p. 105) stated that he is returning to the first name, and includes B. apiciconus in the synonymy of B. sulcatus . Notably, (1843) makes no reference to the work of (1843), suggesting he has not seen that work. (1849, p. 136) followed (1843) and (1843) in referring only 's 1826, Pl. VIII, figs. 3-4 to B. sulcatus and also included B. apiciconus as a synonym. (1849, p. 136) also reproduced the description given by (1843, p. 105). (1849) provided an illustrated guide of fossils of Great Britain and Ireland and listed Belemnites sulcatus from the "Inferior Oolite, Daudry {sic Dundry}, Somersetshire". On his Pl. XXIX, figs. 9-11, he reproduced the illustrations of both specimens of Belemnites sulcatus as given by (1826). By 1854, acknowledged the 1843 work by (p. vi) and, on p. 301, restricted Belemnites sulcatus to 's Pl. VIII, fig. 3 and included B. apiciconus as a synonym. According to article 74.5, none of these publications (, 1843, 1854; , 1843; , 1849; , 1849) qualify as a valid designation of a lectotype for Belemnites sulcatus .
At this point it is worthy of note that Dundry undoubtedly refers to Dundry Hill which exposes an outlier of Aalenian-Bajocian limestone attributed to the Inferior Oolite ( in et al., 1980) and contains abundant belemnites belonging to the genera Belemnopsis , 1878 (inclusive of Belemnites apiciconus ), Holcobelus , 1927, and Brevibelus , 1992 (author's pers. observ., 1992). Oxford is built on the Oxford Clay which here yields common specimens of belemnites referable to Belemnites sulcatus (B. altdorfensis and B. beaumontianus) in the sense of 1870 (, 1870, p. 117).
Neither (1843, 1854) nor (1849) specifically mentioned 's 1826 specimen shown in his Pl. VIII, fig. 5. (1849, p. 140) did describe Belemnites Altdorfensis from the Oxford Clay, and (1854, p. 300) listed Belemnites Beaumontianus from the Oxford Clay of Loch Staffin on the authority of I. . For the Loch Staffin occurrence, (1852, p. lxxvi) reported that Prof. had found a bed with "Ammonites cordatus, Belemnites Owenii, and B. Beaumontianus, [that] distinctly proved it to be of the period of the Oxford Clay".
The available evidence therefore indicates that by the middle of the nineteenth century the name Belemnites sulcatus had first been (tentatively?) restricted to the Oxford Clay form by (1829). Yet, by the 1840s, Belemnites sulcatus was universally used for the belemnite from the Inferior Oolite (the Belemnites apiciconus of ) of southern England and France, whereas the Oxford Clay form was referred to B. altdorfensis or B. beaumontianus .
John ' (1865, 1869, 1870) work on the British Belemnitidae is still a major work for the Jurassic belemnites of England. In revising Belemnites sulcatus, clearly looked towards his earlier record of Belemnites sulcatus in the Kelloways Rock and Oxford Clay of Yorkshire, and not the works of (1843, 1854), (1843), nor (1849). (1870, p. 114) therefore considered that both B. Altdorfensis and B. Beaumontianus were closely related to B. sulcatus and that B. apiciconus was distinct. (1999) suggested that (1870) was the "First Reviser" of Belemnites sulcatus (although even in his monograph, first mentioned Belemnites sulcatus in 1865, p. 5), but had indicated his interpretation of the fossil in 1829. For (1999) therefore, (1870, p. 114), as the First Reviser, restricted B. sulcatus to the form figured by (1826), Pl. VIII, fig. 5. Furthermore, (1869, p. 101) stated that 's 1826, Pl. VIII, figs. 3-4 appeared to represent Belemnites apiciconus , 1827, but did not include it in the synonymy list for B. apiciconus in his monograph. ' 1829 work is hardly a revision of the species, but the works of (1843) and (1843) clearly are and both retain the name Belemnites sulcatus for the form figured as Belemnites apiciconus by (1827). From article 24.2 it is possible to argue that either (1843) or (1843) were first revisers, but not (1870) as indicated by (1999).
The question of whether there has been a valid designation of a lectotype for Belemnites sulcatus is therefore very critical to the understanding of the species concept and its relationship to higher taxonomic rankings. There is no formal nomination of a lectotype in either (1829, 1865, 1869, 1870), (1843, 1854), (1843) or (1849). (1999) took ' 1870 restriction of Belemnites sulcatus to 's 1826, Pl. VIII, fig. 5 as a designation of lectotype, but this is not admissible under article 74.5.
In 1999 (p. 66), formally designated lectotypes for Belemnites altdorfensis and Belemnites apiciconus and these are valid under article 74.5. For Belemnites sulcatus , (1999) did not formally designate a lectotype, but figured "the" lectotype that he considered had been designated by (1870). Since (1870) had never selected a lectotype for Belemnites sulcatus , 's 1999 illustration of a lectotype (on the authority of ), without formal designation, may or may not be a suitable designation of a lectotype according to article 74.5. 's 1999 p. 64 statement in regard to his fig. 6 states "'Inferior Oolite' [=Oxford Clay Formation], ?Dundry near Oxford, England, Lectotype of Belemnites sulcatus" serves as a more formal designation of a lectotype according to article 74.5. In order to stabilize nomenclature, this specimen (, 1826, Pl. VIII, fig. 6) is formally designated lectotype herein in case 's 1999 designation should subsequently prove to be in doubt.
(1999) stated that Belemnopsis in , 1849, was a senior homonym of Belemnopsis , 1878, and attributed the discovery of this homonymy to (1926, p. 65). If correct, this means that Belemnopsis is invalid. The nomenclature issues that would prevail if Belemnopsis in (1849) is deemed valid are far reaching. The generic name Belemnopsis would have to replace Belemnosis in the Family Belemnoseidae , 1922. The relationship and validity of Belemnopsis in , 1849, and Belemnopsis , 1878, therefore, need to be more fully evaluated from insight from further studies of the literature and in light of the rules of the (1999).
In his Catalogue of the Mollusca in the British Museum, (1849) described the species of mollusc then known from the British Isles. In that volume he described the belemnites, some of which have been discussed above, together with various teuthids and sepiids. Amongst the sepiids was J. de C. 's 1829 species Beloptera anomalus which was described by (1849) on page 118. In the section on "Additions and Corrections" at the end of this work, (1849, p. 157-158) added a description of Belemnopsis anomala with the genus attributed to J.E. ', as then unpublished, work on the cephalopods of the London Clay.
' 1849 monograph on the cephalopods of the lower Tertiaries describing a new sepiid taxon (both genus and species) Belemnosis plicata appeared in the Palaeontographical Society volume for the year 1848, which was published in July 1849 ( & , 1877). introduced the new genus Belemnosis, but also introduced a new specific name plicata which, since there was only but a single specimen that had received the specific name anomalus by J. de C. , becomes a junior synonym. (1849, footnote at the bottom of p. 38) indicated the derivation of the name Belemnosis as being from: Βελεμνον, telum and ενωηιs, conjunctio, relating to the transition between belemnites and sepiids. In contrast (1999, p. 60) suggested the suffix –opsis, as used in Belemnopsis in , 1849, was derived from the Greek for 'form' or 'shape' which does not agree with the eponym given by (1849). The question then is: which, Belemnosis or Belemnopsis, has priority?
Dating the publication of 's various catalogues of animals in the British Museum is difficult as each only has the year of publication indicated. (1926) attempted to determine the date of issue of each catalogue (that is issue to book sellers), but could only determine the dates when the Catalogues were laid down on the table for the trustees of the British Museum, and not the actual date when the Catalogues were issued to dealers. (1926) determined that 's Catalogue for 1849 was laid on the table for trustees on the 30th June 1849, and this was confirmed by (1979). If it was immediately submitted to dealers, then 's 1849 work would be deemed to have appeared one month earlier than ' 1849 work (, 1999, article 21.3.1). Yet (1849, p. 158) cited the page numbers from ' 1849 work for the description of both the genus and the species of Belemnopsis plicata as well as giving the plate and figure numbers of the illustration (although he cited the title of the work incorrectly as "Cephalopes of London Clay"). (1849) also amended the specific name from plicata, which must be a junior synonym as there was only one specimen, to anomalus. The citation of actual page and figure numbers, as well as specific names, indicates that (1849) must have seen page proofs or a preprint of 's 1849 work before publication so as to allow him to include these details in his own work. Furthermore, ' 1849 paper was included in the volume of the Palaeontological Society for 1848, suggesting it was completed in 1848 and was awaiting publication; additionally there is no mention of 's 1849 work in (1849). It seems therefore irrefutable that had access to ' 1849 work in the latter stages of completing his own (1849) Catalogue, but that his Catalogue must be deemed to have appeared a month earlier that ' work. According to the Code (, 1999, article 21.8), prior to the year 2000, the distribution of preprints of a work before the recognized publication of the work advances the date of publication, but this does not apply to page proofs, and since no preprints have been recorded it is most likely that saw the page proofs of ' work. Given the fact that acknowledged ' work, it is clear that was not trying to claim authorship of ' generic name and it would also seem clear that Belemnopsis in , 1849, is a spelling mistake for Belemnosis , 1849, even though it was published earlier. It is worthy of note that, according to (1913), there are many spelling mistakes in 's works during the interval 1838-1845, and it would appear that the spelling Belemnopsis is another case.
There are two ways of treating this situation. Firstly, that Belemnopsis is an "incorrect original spelling" of Belemnosis (, 1999, article 32). Secondly, that Belemnosis is an "incorrect subsequent spelling" of Belemnopsis (, 1999, article 33). From the derivation of the name Belemnosis as given by (1849, footnote at the bottom of p. 38) it is clear that Belemnosis was the intended spelling for the generic name and that Belemnopsis is therefore an "incorrect original spelling", rather than an "incorrect subsequent spelling" because 's work appear before that of '. Because (1849) clearly referred to ' 1849 work, this could be taken as an indication of where to find the correct spelling of the generic name as required in article 32.5.1. However this does not entirely fit the article. If we look at current usage, Belemnopsis in , 1849, is not used even by et al. (1998, p. 312; even though on p. 253, Pachybelemnopsis is used in preference to Belemnopsis ) and seems to be universally regarded as an "incorrect original spelling" (e.g., , 1852, p. 17). It is therefore acceptable to consider Belemnopsis 's in (1849) as an "'incorrect original spelling" which therefore cannot enter into homonymy (, 1999, 32.4).
In 1878, introduced a new genus Belemnopsis for a group of belemnites carrying an elongate ventral groove, yet only the volume illustrating the plates, but not the text volume, was issued. Furthermore, there was no indication of a selection of a type species for Belemnopsis (or the other genera). In his volume, (1878) illustrated four species which he included in Belemnopsis, namely: Belemnopsis Altdorfensis (, 1878, Pl. XXIX, figs. 3-4); Belemnopsis Bessina (, 1878, Pl. XXX, fig. 1); Belemnopsis unicanaliculata (, 1878, Pl. XXX, fig. 2, 5); and Belemnopsis sulcata (, 1878, Pl. XXX, figs. 3-4).
The following year, (1882) writing in The Geological Record for 1878 listed the new belemnite genera that (1878) had introduced and recorded that Belemnopsis was preoccupied. (1999) stated that (1926) was the first to recognize that Belemnopsis , 1878, was preoccupied, yet in 1882 clearly indicated that Belemnopsis , 1878, was preoccupied, but without specifying that it was preoccupied by Belemnopsis , 1849. Yet, because Belemnopsis in (1849) must be regarded as an "incorrect original spelling", Belemnopsis , 1878, is an available, and therefore valid, name.
The first designation of a type species for Belemnopsis was made by (1879, p. 91) who selected Belemnites sulcatus as type species. 's 1878 four "species" which he placed in Belemnopsis would now be placed amongst several genera and species, but it is important to record that 's 1878, Pl. XXX, fig. 4, represents Belemnites sulcatus as represented in 's 1826, Pl. VIII, fig. 3 and also equivalent to Belemnites apiciconus of , 1827. (1878) clearly followed (1843, 1854) and (1843) in his concept of B. sulcatus and did not use the name Belemnites apiciconus for this form. Therefore, 's 1879 designation of Belemnites sulcatus as type species for Belemnopsis is valid, contrary to the suggestion by (1999), and did not establish a new nominal species "Belemnopsis sulcatus 1878".
(1878) figured two specimens under the name Belemnopsis sulcata, his plate 30, figure 3 is attributable to Holcobelus subblainvillei () and his plate 30, figure 4 is attributable to Belemnites sulcatus as figured by (1826) plate 8, fig. 3 (= Belemnites apiciconus ) (see , 1999, table 1). As such, one of the specimens illustrated by (1878) as Belemnopsis sulcata belongs to the same species as one of the specimens in the type species of Belemnites sulcatus of (1826). Therefore, 's 1879 nomination of Belemnites sulcatus as type species of Belemnopsis is valid.
However, if 's 1999 selection of a lectotype for Belemnites sulcatus is valid then the type species for Belemnopsis , 1878, can be either Belemnites sulcatus "the nominal species originally cited as type species" as defined by 's 1999 lectotype (article 70.3.1) or Belemnites apiciconus , 1827, "the taxonomic species actually involved in the misidentification" (article 70.3.2). 's 1879 designation of a lectotype needs to be validated by citation to articles 11.10, 67.13 and 69.2.4 to serve stability and universality. The following courses are available: A) to select Belemnites sulcatus as defined by its lectotype (, 1999) as type species of Belemnopsis; or B) to define Belemnites apiciconus , the misidentified species in (1878), as type species of Belemnopsis. Stability is best served by the latter course, and herein Belemnites apiciconus , the misidentified species attributed to Belemnites sulcatus by (1879) and figured as Belemnopsis sulcata by (1878) is selected as type species of Belemnopsis as validated by article 70.3.2.
This paper has discussed the complex nomenclature and literature relating to the species Belemnites sulcatus and has come to the following conclusions.
1. To recognise three specimens as the syntypes of 's Belemnites sulcatus: specimen 1: 's Pl. VIII, figs. 3-4; specimen 2, 's Pl. VIII, fig. 5; and specimen 3, 's 1677, Pl. III, fig. 6. All the specimens appear to be lost or destroyed.
2. To conclude that there was no valid designation of a lectotype under the rules of ICZN for Belemnites sulcatus by (1827), (1829, 1865, 1869, 1870), (1843, 1854), or (1843).
3. To accept 's 1999 nomination of a lectotype of Belemnites sulcatus as 's 1826, Pl. VIII, fig. 5, either directly or by designation herein if such a nomination is not valid according to articles.
4. To accept Belemnopsis in , 1849, as an "incorrect original spelling" of Belemnosis , 1849, as indicated by universal usage. As such Belemnopsis cannot enter into homonymy.
5. To designate Belemnites apiciconus , 1827, as the type species of Belemnopsis , 1878, the species misidentified by , 1878, as Belemnites sulcatus (as subsequently fixed by nomination of a lectotype) which was subsequently nominated as type species of Belemnopsis by (1879).
This action has the advantage of: A) stabilizing the generic name Belemnopsis for a group of belemnites centred around Belemnites apiciconus with a correctly identified type species as illustrated by (1878) and designated by (1879); B) Stabilizing the name Belemnosis , 1849, for the sepiid Belemnosis anomala (J. de C. ); and C) maintaining the previous used family names (Belemnopsiedae , Belemnoseidae ) that are well entrenched in the literature.
No formal descriptions are given here, only systematic lists and brief discussions. For detailed descriptions of taxa see the indicated resources below.
Order BELEMNITIDA , 1895
Suborder BELEMNITINA , 1895
Family HOLCOBELIDAE , 1977
Genus Holcobelus , 1927
Type species. Belemnites munieri , 1878.
Holcobelus blainvillii (, 1830)
1830 Belemnites blainvillii: , p. 37, Pl. 1, fig. 9.
1878 Belemnites unicanaliculatus ; , p. XXX, fig. 5.
Holcobelus munieri (, 1878)
1878 Belemnites munieri: , p. 63, Pl. V, figs. 3-6, 12-14, Pl. VI, figs. 5-11.
1878 Belemnopsis unicanaliculatus ; , p. XXX, fig. 2.
?1878 Belemnites sulcatus ; , p. XXX, fig. 3.
Discussion. et al. (2012) have revised the belemnite Family Holcobelidae , but I disagree that it should be placed in the Belemnopseinae (Pachybelemnopseinae ) and instead place it in the Belemnitinae following (1965).
Family CYLINDROTEUTHIDIDAE , 1919
Subfamily LAGONIBELINAE , 1977
Genus Holcobeloides , 1958
Type species. Belemnites beaumontianus , 1843 (=Holcobeloides altdorfensis , 1827 = Belemnites sulcatus , 1826).
Holcobeloides sulcatus , 1826
1826 Belemnites sulcatus: , p. 59, Pl. VIII, fig. 5 [non figs. 3-4 = Belemnopsis apiciconus (, 1827)]
1827 Belemnites Altdorfensis: , p. 67-69, Pl. 2, fig. 1.
1843 Belemnites Beaumontianus: , p. 118, Pl. XVI, figs. 7, 11.
1849 Belemnites sulcatus ; , p. 248, Pl. XXIX, fig. 10 [non fig. 9 = Belemnopsis apiciconus (, 1827)]
1870 Belemnites sulcatus ; , 115-117, Pl. XXIX, figs. 71-73, Pl. XXX, figs. 74-75.
1878 Belemnopsis altdorfensis ; , Pl. XXIX, figs. 3-4.
Discussion. The genera of the Family Cylindroteuthididae have been revised recently by (2011).
Suborder BELEMNOPSEIDINA , 1965
(=Pachybelemnopseina in et al., 1998)
Family BELEMNOPSEIDAE , 1922, emend , 1946
(=Mesohibolitidae , 1983)
Genus Belemnopsis , 1878
Type species. Belemnites apiciconus , 1827 (= Belemnites sulcatus as designated type species by , 1879, and figured as Belemnopsis sulcatus by , 1878).
(= Pachybelemnopsis , 1980, type species: Belemnites canaliculatus , 1820; non Belemnopsis , 1849, which is an unavailable name which cannot enter in homonymy because it is based on an incorrect original spelling of Belemnosis , 1849)
Belemnopsis apiciconus , 1827
1826 Belemnites sulcatus: , p. 59, Pl. VIII, figs. 3-4 [non fig. 5 = Holcobeloides sulcatus (, 1826)]
1827 Belemnites apiciconus: , p. 69, Pl. 2, fig. 2.
1843 Belemnites sulcatus ; , p. 177 (restricted to 's 1826, Pl. VIII, fig. 3).
1849 Belemnites sulcatus ; , p. 136 (restricted to 's 1826, Pl. VIII, figs. 3-4).
1849 Belemnites sulcatus ; , p. 248, Pl. XXIX, fig. 9 [non fig. 10 = Holcobeloides sulcatus (, 1826)]
1854 Belemnites sulcatus ; , p. 301 (restricted to 's 1826, Pl. VIII, fig. 3).
1869 Belemnites apiciconus ; , p. 101-102, Pl. XXV, fig. 58.
Type specimen. Specimen figured by , 1827, Pl. 2, figs. 2, 2a; designated lectotype by (1999, p. 66).
Order SPIRULIDA , 1912
Family BELEMNOSEIDAE , 1869
Genus Belemnosis , 1849
(= Belemnopsis in , 1849, an unavailable incorrect original spelling of Belemnosis , 1849)
Type species. Beloptera anomalus J. de C. , 1829.
Belemnosis anomala (J. de C. , 1829)
1829 Beloptera anomalus J. de C. , p. 183, Pl. 591, fig. 2.
1838 Beloptera anomala ; , p. 43, Pl. XXIX, figs. 23-24.
1849 Beloptera anomala : , p. 118.
1849 Belemnopsis anomala (): , p. 157-158.
1849 Belemnosis plicata: , p. 40, Pl. 2, fig. 3a-c.
I would like to thank the reviewers for their comments on this paper, especially Philippe for taking time to go through my arguments and making detailed suggestion with regard to the operation of the ICZN rules. Nico M.M. is thanked for correcting some problems with the paper.
E. (1878).- Atlas. Première partie. Fossiles principaux des terrains.- Explication de la Carte géologique de la France, Paris, t. IV, pt. 1, 158 Pls.
URL: http://gallica.bnf.fr/ark:/12148/bpt6k6310840g/f11.image.r=
M.H. D. de (1927).- Mémoire sur les Bélemnites, considérées zoologiquement et géologiquement.- F.G. Levrault, Paris, 136 p. (5 Pls.).
URL: https://archive.org/stream/mmoiresurlesble00blaigoog#page/n13/mode/2up
J.F. (1882).- , Prof. E. (p. 315). In: 2. Invertebrata.- The Geological Record for 1878, London, p. 313-350.
URL: https://archive.org/stream/geologicalrecor04unkngoog#page/n350/mode/2up
T. (1838).- Genus XI.- Belemnites.- .- Illustrations of the fossil conchology of Great Britain and Ireland, with descriptions and localities of all the species hiterto discovered.- Smith, Elder & Co., London, nº 8, Pl. XXIX; nº 10, p. 41-43.
URLs: https://archive.org/stream/illustrationsoff00brow#page/n121/mode/2up and https://archive.org/stream/illustrationsoff00brow#page/41/mode/2up
T. (1849).- Illustrations of the fossil conchology of Great Britain and Ireland, with descriptions and localities of all the species.- Smith, Elder & Co., London, Concluding part, nº 29-35, p. 137-273.
URL: https://archive.org/stream/illustrationsoff00brow#page/n437/mode/2up
A.B. & D.C.H. (2007).- New Middle and Upper Jurassic belemnite assemblages from West Antarctica (Latady Group, Ellsworth Land): taxonomy and paleobiogeography.- Palaeontologia Electronica, vol. 10, Issue 1 (6A), 29 p.
J.C.W., K.L., C.F., H.S., W.A. & J.K. (1980).- A correlation of Jurassic rocks in the British Isles: Part two: Middle and Upper Jurassic.- Geological Society of London, Special Report, nº 15, 109 p.
M. (1879).- Atlas du IVe volume de l'Explication de la Carte géologique de la France.- Bulletin de la Société géologique de France, Paris, (3ème série), t. 7 (1878-1879), p. 91-92.
URL: http://jubilotheque.upmc.fr/img-viewer/fonds-bulsgf/GB_000051_001/Contenu/JPEG_HD/viewer.html?ns=GB_000051_001_J3_0001.jpg
P. (1992).- The British Toarcian (Lower Jurassic) belemnites. Part 2.- Monograph of the Palaeontographical Society of London, no. 144 (1991), p. 50-79.
O.S. (2011).- Subfamily classification within the Cylindroteuthididae (Belemnitida).- News of palaeontology and stratigraphy, vol. 16-17; Supplement to Russian Geology and Geophysics, t. 52, p. 103-108. [in Russian]
F.E. (1849).- A monograph of the Eocene Mollusca, or descriptions of shells from the older Tertiaries of England. Part I. Cephalopoda.- Palæontographical Society Monograph, London, p. 1-56 (Pls. I-IX).
URL: https://archive.org/stream/monographofeocen02edwa#page/n7/mode/2up
F.E. & S.V. (1877).- A monograph of the Eocene Mollusca, or, descriptions of shells from the older Tertiaries of England. Part IV. Pulmonata and Prosobranchiata.- Palaeontographical Society Monograph, London, p. 331-361 (Pl. XXXIV)
URL: https://archive.org/stream/monographofeocen02edwa#page/n17/mode/2up
E. (1878).- Le Jura Normand. 2ème Livraison. Monographies VI. Assises supérieures des marnes infra-oolithiques.- Savy, Paris, 78 p. (XI Pls.).
J.E. (1849).- Catalogue of the Mollusca in the collection of the British Museum. Part 1: Cephalopoda Antepedia.- Spottiswoodes and Shaw, London, 164 p.
URL: https://archive.org/stream/cataloguemollus01zoolgoog#page/n6/mode/2up
V.A. (1958).- New Upper Jurassic belemnites from the Russian Platform.- Bjulleten Moskovskogo Obshchestva Ispytateley Prirody, (Seriya Geologicheskikh), Moscow, vol. 33, nº 4, p. 158-159 [in Russian].
V.A. (1977).- On a revision of Jurassic belemnites.- Bjulleten Moskovskogo Obshchestva Ispytateley Prirody, (Seriya Geologicheskikh), Moscow, vol. 52, nº 2, p. 103-117 [in Russian].
A.N. (1852).- Indicis generum malacozoorum supplementa et corrigenda.- Theodori Fischer, Cassellis (Kassel), 140 p.
W. (1852).- Anniversary address of the president.- The Quarterly Journal of the Geological Society of London, vol. 8, p. xxi-lxxx.
URL: https://archive.org/stream/quarterlyjournal81852geol#page/n33/mode/2up
(1999).- International Code of Zoological Nomenclature.- 4th Edition, London, 306 p.
URL: http://www.nhm.ac.uk/hosted-sites/iczn/code/
T. (1913).- A collation of the molluscan parts of the synopses of the contents of the British Museum, 1838-1845.- Proceedings of the Malacological Society of London, vol. X, p. 294-309.
URL: https://archive.org/stream/proceedingsofmal21041912213mala#page/294/mode/2up
J.A. (1946).- Zur Kenntnis der oberkretazischen Belemniten.- Geologiska Föreningens i Stockholm Förhandlingar, vol. 68, p. 87-105.
J.A. (1965).- Taxonomy and phylogeny of fossil Coleoidea (= Dibranchiata).- Papers of the Geological Survey of Canada, Ottawa, vol. 65, nº 2, p. 72-76.
A.R. (1989).- The " Catalogues". [Mollusca] of the British Museum.- The Nautilus, Sanibel, vol. 103, nº 3, p. 113–115.
J.S. (1826).- V. - Observations on belemnites.- Transactions of the Geological Society of London, (Series 2), vol. 2, p. 45-62 (Pls. VII-IX).
J. (1843).- A catalogue of British Fossils comprising the genera and species hitherto described; with references to their geological distribution and to the localities in which they have been found.- John Van Voorst, London, 1st Edition, 222 p.
J. (1854).- A catalogue of British Fossils: comprising the genera and species hitherto described; with references to their geological distribution and to the localities in which they have been found.- Printed privately, London, 2nd Edition, 372 p.
URL: https://archive.org/stream/acataloguebriti00morrgoog#page/n4/mode/2up
A. (1922).- Die fossilen Tintenfische. Eine paläozoologische Monographie.- G. Fischer, Jena, 322 p.
URL: http://www.biodiversitylibrary.org/item/18349#page/5/mode/1up
V.M. (1983).- Early Cretaceous belemnites from the south of the USSR. In: Y.I. & K.N., eds., Systematics and ecology of cephalopods.- Zoological Institute, Leningrad, p. 42-43 [in Russian].
A. d' (1843).- Terrains oolitiques ou jurassiques. In: Paléontologie française. Description zoologique et géologique de tous les animaux mollusques et rayonnés fossiles de France, comprenant leur application à la reconnaissance des couches.- Tome Premier, Cosson, Paris, 642 p.
URL: https://archive.org/stream/palontologiefr11orbi#page/n5/mode/2up
J. (1829).- Illustrations of the geology of Yorkshire; or, A description of the strata and organic remains of the Yorkshire coast: accompanied by a geological map, sections, and plates of the fossil plants and animals, Part 1, The Yorkshire coast.- Privately printed, York, 1st Edition, 192 p. (XIV Pls.)
URL: http://www.biodiversitylibrary.org/item/74325#page/9/mode/1up
J. (1835).- Illustrations of the geology of Yorkshire; or, A description of the strata and organic remains of the Yorkshire coast: accompanied by a geological map, sections, and plates of the fossil plants and animals, Part 1, The Yorkshire coast.- John Murray, London, 2nd Edition, 184 p. (XIV Pls.).
URL: http://books.google.fr/books?id=qm_nAAAAMAAJ&hl=fr&pg=PR1#v=onepage&q&f=false
J. (1865).- A monograph of British Belemnitidæ.- Monographs of the Palaeontological Society of London (1863), vol. XVII, p. 1-28.
URL: https://archive.org/stream/monographof171865pala#page/n251/mode/2up
J. (1869).- A monograph of British Belemnitidæ. Part IV.- Monographs of the Palaeontological Society of London (1868), vol. XXII, p. 87-108 (Pls. XXI-XXVII).
URL: https://archive.org/stream/monographof221869pala#page/n307/mode/2up
J. (1870).- A monograph of British Belemnitidæ. Part V.- Monographs of the Palaeontological Society of London (1869), vol. XXIII, p. 109-128 (Pls. XXVIII-XXXVI).
URL: https://archive.org/stream/publication12britgoog#page/n161/mode/2up
J.F. (1912).- Cephalopoda. Paläontologie.- Handwörterbuch der Naturwissenschaften, Jena, Band 2, p. 265-296f.
URL: http://www.biodiversitylibrary.org/item/17519#page/279/mode/1up
R. (1677).- The natural history of Oxford-fhire, being an effay toward the natural hiftory of England.- Theater, Oxford, 358 p. (XVI Pls.).
URL: http://www.biodiversitylibrary.org/item/64513#page/7/mode/1up
W. (1980).- Revision der Belemniten der Schwäbischen Jura. Part 7.- Palaeontographica Abteilung A, Stuttgart, vol. 169, p. 128-206.
W. (1999).- Taxonomic status of the belemnite genus Belemnopsis 1878.- Palaontologische Zeitschrift, Stuttgart, vol. 73, nº 1-2, p. 59-76.
W., N. & C. (1998).- Cephalopoda dibranchiate fossils (Coloidea) II. In: F., ed., Fossilium Catalogus I: Animalia pars 135.- Backhuys Publishers, Leiden, 519 p.
E.F. von (1820).- Die Petrefaktenkunde auf ihrem jetzigen Standpunkte durch die Beschreibung seiner Sammlung versteinerter und fossiler Überreste des Thier- und Pflazenreichs der Vorwelt erläutert.- Becker, Gotha, 437 p. (XV Pls.).
URL: http://books.google.fr/books?id=4SQ-AAAAcAAJ&hl=fr&pg=PR1#v=onepage&q&f=false
C.D. (1926).- Dates of publication of early catalogues of natural history issued by the British Museum.- Journal of Natural History, (Series 9), vol. 17, nº 98, p. 271-272.
J., continued by J.D.C. (1829).- The mineral conchology of Great Britain; or coloured figures and descriptions of these remains of testaceous animals or shells, which have been preserved at various times and depths in the Earth.- Richard Taylor, London, vol. VI, 250 p.
URL: http://www.biodiversitylibrary.org/item/50339#page/5/mode/1up
E. (1919).- Die Systematik der Belemniten.- Jahresbericht des Niedersächsischen Geologischen Vereins, Hannover, vol. XI, p. 1–59.
E. (1927).- Zur Systematik und Stratigraphie median gefurchter Belemniten.- Jahresbericht des Niedersächsischen Geologischen Vereins, Hannover, vol. XX, p. 112-136.
E. (1926).- Miscellanea nomenclatorica zologica et palaeontogica I-III.- Archiv für Naturgeschichte, Berlin, (Abeitlung A) Band 92, Heft 8, p. 30-75.
P.L. (1830).- Observations sur les bélemnites.- Mémoires de la Société d'histoire naturelle de Strasbourg, t. I, p. 1-70 (VIII Pls.).
URL: https://archive.org/stream/mmoiresdelaso01soci#page/n13/mode/2up
R., N. & W. (2012).- The belemnite Family Holcobelidae (Coleoidea) in the European Jurassic: systematics, biostratigraphy, palaeobiogeography and evolutionary trends.- Palaeodiversity, Stuttgart, vol. 5, p. 13–49.
T. (1869).- On the chief groups of the Cephalopoda (A paper read before the Geologists' Association, Nov. 1st, 1867.).- Proceedings of the Geologists' Association, London, Supplement, vol. I (1864-1871), p. 181-206.
K.A. von (1895).- Grudzüge der Paläontologie (Paläozoologie). 1 Abeitlung: Invertebrata.- R. Oldenbourg, München & Berlin, viii + 733 p. (Dibranchiata, p. 468-481).
URL: https://archive.org/stream/grundzgederpal00zitt#page/468/mode/1up