AMERICAN MUSEUM NOVITATES Published by Number 1314 THE AMERICAN MUSEUM OF NATURAL HISTORY March 14, 1946 New York City

NOTES ON STROMBUS DENTATUS LINNE AND THE STROMBUS URCEUS COMPLEX BY HENRY DODGE

A remark made by Tryon in his mono- latus Schumacher, 1817. This complex graph on the Strombidae (1885, p. 118) in presents such extremes of form, however, discussing Strombus dentatus Linn6 has that further study may justify its separa- prompted me to examine the taxonomic tion into subspecies or even species. history of the species in the so-called den- tatus group. Tryon there said: "The 1 difference between this species and S. On the first point we should turn im- urceus is so slight, and there is so much mediately to the Linnaean descriptions: variation in the shells, that it is very Strombus urceus LINNAEUS, 1758, p. 745, No. doubtful whether can be 440; 1767, p. 1212, No. 512. their separation "S. testae [sic] labro attenuato retuso brevi maintained." striato, ventre spiraque plicato-nodosis, apertura A reading of the meager references to bilabiata inermi." this group, a study of the synonyms in- TRANSLATION: Shells with a "thinned-out," of a reflected, short, and ridged lip, body-whorl and volved, and an examination consider- spire plicate-nodose, aperture bilabiate and lack- able series of specimens disclose an ob- ing armature. vious confusion which appeared as early as Strombus dentatu8 LINNAEUS, 1758, p. 745, Gmelin and has persisted in the minds of No. "o"; 1767, p. 1213, No. 513. I have "S. testa labro attenuato brevi dentato, ventre all but a few authors since his day. spiraque plicatis." concluded that: TRANSLATION: Shell with a "thinned-out," 1. Dentatus Linne is not identical with short and toothed lip, body-whorl and spire urceus Linn6 or with any form of the urceus plicate. of authors, and Tryon's remark is true The figures shown in the synonymy of only if we assume that he was speaking of Linne's urceus all show approximations to the shells which are today almost univer- the more or less strongly plicated form of sally known as dentatus and urceus Linn6 the urceus of authors, although it is to be in our collections. It is not true if he noted that this is not the form usually meant the dentatus and urceus described labeled urceus in our collections, which is by Linnaeus. in most cases the larger, more graceful, and 2. Dentatus as described in the "Sys- less plicate shell. It is unfortunate that tema naturae" is identical with the shell Linnaeus did not supply a synonymy for described by Gmelin in 1791 and Lamarck dentatus in either the tenth or twelfth in 1822 as tridentatus, by Chemnitz in 1777 edition of the "Systema." Had he done as dentatus, and by Reeve in 1851 as sana- so the question of its identity might never rensis. The other names mentioned should have arisen. therefore be thrown into the synonymy of It is obvious that the confusion in the dentatus Linne and that shell disassociated identity of dentatus and its fancied relation from any form of urceus Linn6. to urceus have arisen from a careless reading 3. Urceus Linn6 is tentatively to be of the Latin description. But the con- listed as unidentifiable. The various forms clusion seems inescapable that Linnaeus of the complex known as the urceus of was describing two entirely different shells. authors seem to be one species for which The reflection of the lip and the presence the earliest valid name is Strombus ustu- of nodes in urceus are not indicated in the 2 AMERICAN MUSEUM NOVITATES [No. 1314

I 2

3 4

Fig. 1. Strombus ustulatus Schumacher, form laevis, new name. (S. urceus of most authors, non Linne.) No locality data. A.M.N.H. No. 12927. Fig. 2. Strombus dentatus Linn6 (S. tridentatus Gmelin). Philippine Islands. Fig. 3. Strombus ustulatus Schumacher, form plicatus, new combination. (S. dentatus of most authors, non Linn6.) Catanduanes, Philippine Islands. Fig. 4. Strombus floridus Lamarck (S. mutabilis Swainson). "Indian Ocean." A.M.N.H. No. 49418. Figures 1 and 3 are examples of the extreme forms of S. ustulatus from the standpoint of sculpture. All figures X 1. Photographs by Marion A, Bills. 1946] STROMBUS DENTATUS AND THE S. URCEUS COMPLEX 3

5 6 7 8 Back views of shells illustrated in figures 1 to 4. Fig. 5. Strombus ustulatus Schumacher, form laevis. Fig. 6. Strombus dentatus Linn6. Fig. 7. Strombus ustulatus Schumacher, form plicatus. Fig. 8. Strombusfloridus Lamarck. All figures X 1. Photographs by Marion A. Bills. description of dentatus. The aperture of the lip. In the latter they are true teeth urceus is stated to be "inermi," whereas on the lip. the lip of dentatus is "dentato." In the There may have been a difference of latter species he was describing a shell with opinion as to whether the word "brevi" in a toothed lip. Linnaeus' language can be the description of both urceus and dentatus read in no other way. It is true that in is an adjective modifying "labro" or an ad- certain other descriptions Linnaeus digni- verb modifying "dentato." I suggest fied the striae or furrows of the inner aspect that a study of the descriptions on page of the lip (which are present in all shells of 745 of the tenth edition of the "Systema," the group here discussed) with some name which contains two broad-lipped species, derived from the Latin "dens." I have, gigas and latissimus, and the several short- however, examined the descriptions of lipped species, epidromis, canarium, vitta- every other species of gastropod for which tus, urceus, and dentatus, should satisfy he used "labro dentato" or words of like one that "brevi" is an adjective modifying import, and can report that in no case did "labro." It must be admitted that Lin- he employ such an expression for a shell naeus used "brevi" very broadly as applied which did not have true teeth on the edge to the lips of the species mentioned. It is of the lip or on its inner aspect. As an true that they are all short-lipped, but the illustration, compare Voluta mercatoria shortness varies in degree. The true denta- (1758, No. 357)-our Pyrene mercatoria- tus Linn6 (tridentatus Gmelin) has an un- ("labroque introrsum gibbo denticulato") usually abbreviated lip, and Linnaeus with Murex fusus (1758, No. 478)-our might well have indicated this by more Tibia fusus-("labro dentato"). In the emphatic phraseology. first case the "teeth" are mere ridges inside Thus dentatus possesses a characteristic 4 AMERICAN MUSEUM NOVITATES [No. 1314 which is not present in any of the forms of majority of specimens labeled urceus have the urceus complex, unless one is willing to a colorless or yellow-orange columella and use the word "teeth" for the irregularities lip. The black is the exception. The or crenulations produced by the stromboid black individuals were mostly of the notch. These irregularities, which are smoother form in the series examined, well shown in the figure from Chenu and although the plicated form showed a few Reeve reproduced by Tryon in volume 7 of black specimens. the "Manual of conchology," are often In all these variations I failed to find, pronounced, but in no case are there more between lots of individuals possessing a than two, and these are short, blunt, and given group of traits and other lots possess- rounded and are mere modifications of the ing other groups of traits, any real points of crenulations present in most forms of the cleavage sufficiently marked to justify the urceus complex. If these features may be conviction that any form is entitled to called "teeth," then virtually every Strom- specific or subspecific rank. The varia- bus would have to be described as "den- tions, although presenting very wide ex- tato." tremes, are nevertheless connected by such I have examined a considerable series of a complete chain of intermediates that I specimens labeled urceus Linn6, dentatus cannot find any evidence of discontinuity Linn6, and other names which will be at any point. Nor have I been able to mentioned later, in several private collec- formulate any conclusion as to geographical tions and in the collections of two large races upon the stated locality of the series museums, and in every case those identified examined. The most extreme forms are as dentatus were, within the limits of the found over the entire range of the shells in- variations present in this complex, very volved, which extended from Mindoro in close to those identified as urceus. I am the Philippines to Australia, the New therefore forced to the conclusion that both Hebrides, and the Paumotus, and as far the dentatus and the urceus of authors and west as the Red Sea and Mauritius. Nor of our collections are very closely allied if were any of the lots examined sufficiently not conspecific. documented to enable one to determine The noticeably variable traits are: the whether temperature or salinity of water, length, prominence, and spacing of the type of bottom, nature of food, or any axial plications on the body whorl; the other ecological factors could be given any degree to which the stromboid notch is de- weight. There is, perhaps, room for veloped; the degree and form of the further useful study along this road. thickening of the lip; the sculpture of the spire and the color pattern. The most im- 2 portant color variation is the presence of a If dentatus Linn6 was not the dentatus black or purplish black columella or outer of our museums, what was it? lip or both. This color is so striking that Chemnitz (1788, vol. 10, pp. 220-221), many of the early conchologists mention it in describing dentatus, correctly read the in describing urceus, thus making it a defi- Linnaean description calling it "Die Ge- nite characteristic of the species. (Cf. zahnte Flugelschnecke" and specifically Lamarck, 1822, "labrum caudaque nigri- describing its lip as "labro infra dentato." cante . . . fauce nigra"; Chemnitz, 1845 He cited figures from Seba showing a shell edition, where the shell is called "Die with a definitely toothed lip as distinct Schwartzmundige Flutgelschnecke"; Knorr, from a crenulated lip or one with "teeth" 1768, whocallsit "Die Schwartzmiundchen"; consisting of the sides of the stromboid and Reeve, 1851, who says, vol. 6, pl. 11, notch. The figures shown in the volume of "This species may generally be recognized plates appended to volume 10 of the by the dark purple-black coloring of the "Conchylien Cabinet" are the most ac- columella and aperture, which in some curate pictures of the tridentate Strombus specimens is, however, only partially indi- that have appeared in any iconography cated.") The fact is that the great with the exception of that of Reeve. 1946] STROMBUS DENTATUS AND THE S. URCEUS COMPLEX 5

Chemnitz, however, complicated the issue name (op. cit., p. 210) of which he says in his by describing another shell, S. samar, new French description, "its right border is not name (1788, vol. 10, p. 221). While samar dentate but shows at its lower end the was not described as having a dentate lip, characteristic sinus of the genus." He its pictorial synonymy showed a figure follows this by citing S. dentatus Linn6 as a which probably represents a young tri- synonym, thus becoming the first author dentatus Gmelin with very rudimentary definitely to identify dentatus Linn6 with teeth. Yet the name is in use today, and a member of the urceus complex (urceus of many specimens of typical tridentatus authors). labeled samar Chemnitz are found in our It is not possible within the limits of this collections. I have no idea what Chemnitz paper to discuss the opinion of all the was describing in his samar. Either the authors who have listed this species. name must be rejected as unrecognizable Suffice it to say that Chemnitz and his or be made a synonym of dentatus Linn6 revisers in the 1845 edition, Sowerby (1847, according to the amount of clairvoyance Strombus, p. 31), and Hanley (1855, p. 276) we can exert in reading Chemnitz' descrip- either directly or indirectly properly identi- tion. Sowerby's Strombus bullatus has fied dentatus Linn6 with tridentatus Gmelin, been suggested as being equal to samar, thus correctly interpreting the Linnaean but the vagueness both of Chemnitz' description. All the others, notably figure and of Sowerby's figures of bullatus Gmelin, Lamarck and his revisers, Deshayes precludes any certain identification. and Milne Edwards (1845, vol. 9, p. 706), The mistake which has plagued dentatus Reeve (1851, vol. 6, pl. 9, fig. 17), and Linn6 began with Gmelin. In the thir- Tryon (1885, vol. 7, p. 118) have continued teenth edition of the "Systema naturae" he the old error of identifying it with some listed two shells described as having form of the urceus of authors. dentate lips although only one is known: Hanley was the last writer who properly S. dentatus and S. tridentatus (1791, p. read the Linnaean description. Since his 3519, Nos. 31 and 30, respectively). The day I can find no mention or figure of description of the first is copied bodily dentatus which suggests that it is anything from Linnaeus with a long sub-description else than a form of which adds nothing in regard to the lip. the urceus of authvrs of The description of the second is a good the more plicate type, and the labels in our diagnosis of the tridentate Strombus, the collections follow this lead. If we base our words opinion upon the number of supporters of "labro triacantho" being much more each theory certainly the evidence is pre- graphic than the language used by Lin- ponderantly in naeus. It is impossible to guess what favor of the view presented shell in this paper. And if we study the figures Gmelin thought he was presenting in cited by all the authors through Hanley, his dentatus. The figures he cited for it and particularly if we go to the final and are varied. Some clearly show tridentatus, best and others picture one or another of the source-Linnaeus' own language-we forms of the urceus of authors. Whether are forced to the conclusion that the his lapsus dentatus in our museums is not the dentatus means that he thought that his of Linnaeus. The very name given to the tridentatus was hitherto undescribed or latter was a description of it, and, inci- whether he meant to change the Linnaean dentally, it is the only gastropod to which specific name to his own and then restore he gave it for a form of urceus is only a matter of this name. That fact is surely conjecture. additional evidence. Lamarck was also in error in his concep- tion of dentatus Linn6. He describes 3 tridentatus (1822, vol. 7, p. 209) very I have already said that I cannot separate graphically, using the words "basi tri- out any species or subspecies from among dentato," and supplies a partially correct the very variable group called the "urceus synonymy. He also lists S. plicatus, new of authors," in which I include the dentatus 6 AMERICAN MUSEUM NOVITATES [No. 1314 of most authors. The larger question is: able to suppose that Hanley utilized these Just what shell was Linnaeus describing in works. urceus? The following notes suggest that Watson (1886, p. 417) followed Hanley the urceus of all authors following Linnaeus in holding that the urceus of authors was was a different shell from that described by not the urceus of Linnaeus and also per- him, and that the identity of his urceus has petuated what I feel was Hanley's error in not been satisfactorily determined. believing that S. floridus Lamarck (S. For almost a century after Linnaeus I mutabilis Swainson) was the Linnaean can find no comments in the literature sug- type. In looking for a valid name for the gesting that the urceus of all authors had urceus of authors he chooses the earliest not been described in the "Systema." The post-Linnaean name, Canarium muricatum first who raised the question was Hanley. Martini, 1777, adding that after Martini On page 275 of his "Ipsa Linnaei conchylia" the next in line of priority would be (1855) he says: "One regrets to disturb a Strombus ustulatus (Carnarium ustulatum) long established identification, yet truth Schumacher, 1817. As the Martini names compels me to declare that the S. urceus have been officially rejected as not being of modern writers is not the true representa- consistently binomial, the first valid name is tive of the species thus named by our that of Schumacher. As will appear, I author. We shall not find among the disagree with Watson's identification of S. synonyms a single characteristic drawing of urceus Linn6 with S. floridus Lamarck. that shell . . . On the contrary it is the S. Tryon's description of urceus Linn6 mutabilis (Seba Mus. vol. iii, pl. 60, f, 28 (op. cit., p. 118) recalls the smooth form of etc.) and the S. plicatus (Rumph. pl. 37, the urceus of authors for which I propose f.T. etc.), the former of which is marked for the name: form laevis. As to the color of this species in the Linnaean cabinet, where the aperture he says (loc. cit.), "lip, aperture the supposititious urceus is not present, to and columella deep orange-brown, or chest- which our attention is directed by the nut, or more usually deep chocolate or references." black," showing that he was impressed by As I am treating S. plicatus Lamarck as the "Schwartzmiindige" form as being one of the forms of the urceus of authors, rather typical than otherwise. He makes and as it will be shown that Hanley was in the species synonymous with S. ustulatus error as to the identity of mutabilis Swain- Schumacher, 1817, S. incisus Wood, 1828, son with urceus Linn6, the only helpful part and S. anatellus Duclos (in Chenu), 1844. of his comment is his conviction that the His figures (pl. 6, figs. 65-67) show two post-Linnaean authors were not describing smooth forms and one apparently with no Linnaeus' urceus. One reason given by plications but with strong nodes at the him for this opinion was that "the striking shoulder; two with a strongly plicated color of the aperture in urceus was not spire and one with the spire virtually mentioned in the description" (loc. cit.). smooth. Tryon makes no suggestion that This undoubtedly refers to the black lip there is any difference between the urceus and columella already discussed. Thus of authors and the species that Linnaeus my strong impression is that Hanley meant, described. by the expression "the urceus of authors," C. Hedley (1904, p. 188) also appears to the black-lippe(d form. The two most agree with Hanley that urceus Linne was recent iconographies of the Strombidae in S.floridus Lamarck (S. mutabilis Swainson) Hanley's day were the second edition of the and adds (loc. cit.) "the urceus of authors "Conchylien Cabinet" (1845, vol. 4) and is S. ustulatus Schumacher 1817." Reeve's "Conchologica iconica" (1851, W. Adam and E. Leloup (1938, pp. vol. 6), both of which mention the black 112-116) believed that the urceus of authors aperture as a specific determinant of was identical with S. plicatus Lamarck for urceus Linne. Likewise the best of Sower- which they gave an exhaustive synonymy by's figures in the "Thesaurus conchi- including S. dentatus Gmelin, 1791. They liorum" shows this form. It seems reason- were-unwilling, however, to pass definitely 1946] STROMBUS DENTATUS AND THE S. URCEUS COMPLEX 7

upon the identity of Linnaeus' urceus, but someone following him to whom the custody do disagree with Hanley, Watson, and of his cabinet was entrusted. A reading of Hedley, saying (op. cit., p. 115): "As to the the Introduction to Hanley's work (supra) interpretation of Strombus urceus Linne we should convince one that not only the vicis- do not agree that it is identical with S. situdes which the collection has suffered but floridus Lamarek" (translation). Linnaeus' own inaccuracies as well leave This is the state in which the problem of much room for charges of error of this sort. the identity of urceus Linn6 is left today. I must therefore conclude that urceus Hanley, Watson, and Hedley believe that Linn6 is not floridus Lamarck. It is not he was describing S. floridus Lamarck. the black-lipped form of the urceus of Certainly if that shell is present in the authors as that form also has a smooth or Linnaean cabinet, as Hanley reported, and almost smooth body-whorl. It may have marked for urceus, that fact is weighty but been a shell something like plicatus not conclusive evidence. In any event it Lamarck. But on the state of the evidence would be useful to check Hanley's state- I am content to consider it among the ment by an examination of the collection "lost" species of Linnaeus. de novo as soon as that becomes possible. In summary I suggest: Linnaeus' description of urceus, however, 1. That S. urceus of authors not Linn6, omits many of the most characteristic S. dentatus of authors not Linne, S. plicatus features of floridus. Hanley uses Lin- Lamarck, and the other names cited as naeus' omission of any reference to "the synonyms of urceus and dentatus Linn6 by striking color of the aperture" as evidence Tryon are all forms of a variable species that he was not describing the urceus of which should receive the name Strombus authors. Might it not also be said that ustulatus Schumacher, 1817, at least while the omission of a reference to the rosy pink awaiting further study. I here propose color of the aperture of floridus was evi- for the extremely smooth form of ustulatus dence that he was not describing that shell? the name "form laevis," and for the plicate In fact the description of urceus, although form, the name "form plicatus." broad enough to cover many forms in this 2. That S. urceus Linn6 be rejected as general group, is woefully inapplicable to an undetermined species. floridus, and I will always contend that 3. That S. tridentatus Gmelin, 1791, Linnaeus' descriptions, short though they S. tridentatus Lamarck, 1822, and S. may be, are entitled to more weight than samarensis Reeve, 1851, are all synonyms the figures he cites or the presence or ab- of a well-defined species with extremely sence of marked specimens in his collection. constant characteristics which should re- In addition to the failure to mention the tain the first name of S. dentatus Linn&. pink color of the aperture there is no hint It is unfortunate that the well-established of the characteristic shoulder at the upper name tridentatus should have to be end of the lip. There is no mention of the abandoned in favor of the less graphically folds across the entire length of the colu- descriptive dentatus, particularly in view of mella. Most significant of all, the words the confusion of that name with certain "ventre spiraque plicato-nodosis" in the forms of the urceus of authors. (It is con- description of urceus are utterly inapt as ceivable that the International Commission applied to floridus. on Zoological Nomenclature might hold In the last analysis the only evidence in that Linnaeus' brief and somewhat unen- favor of floridus as the representative of lightening description of dentatus and his urceus Linne is the fact that a specimen of failure to provide any synonymy or locality floridus was said to have been found in would preclude the retention of the Lin- Linnaeus' collection. This may well have naean name, in which case the species been a mistake of Linnaeus himself or of would become S. tridentatus Gmelin, 1791.) 8 AMERICAN MUSEUM NOVITATES [NO. 1314

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