One of the more unusual marine mollusk records for
northeast Florida waters is the occurrence of the Atlantic Partridge Tun. It is
one of a very few locally-collected species on which I personally have never
laid a hand, but documentation was secured by Charlotte Lloyd, who came across a
living specimen at a depth of 128 feet 45 miles east of Mayport on August 6,
1991. Being conservation-minded, yet wanting to placate skeptics of "(shell)fish stories," she chronicled the encounter on videotape just before releasing the
snail!
Aside from Charlotte's unusual coup, there is a bit
more to say about this critter - partly its natural history, but more so the
history of its naming. The unraveling of this paper trail took me through
two and a half centuries of literature and provided a few general lessons in the
sometimes complex process of taxonomic
nomenclature.
The Atlantic Partridge Tun
(see
figure 1., a 40 mm. juvenile from Eleuthera, Bahamas
in the Lee Collection), is widely known as Tonna maculosa (Dillwyn,
1817), a name proposed as "Buccinum perdix .... Variety ... Buccinum
maculosum." on page 583 of volume 2 of this work (see bibliography for this
and other citations). Lewis Weston Dillwyn recorded two references. The first
was "Solander [actually Lightfoot, 1786 ] lot 3050 p. 136" [sic; error
for 137]. In Lightfoot (1786) this entry reads "A very fine variety of Buccinum
maculosum, S. or spotted Tun, undescribed from New Holland -
very rare" [Lightfoot's italics]. The "S." connotes a name in a
Daniel Solander manuscript - possibly only on the label appended to the specimen
in question. Unfortunately the passage I quoted was all Lightfoot wrote; there
is no cited illustration and insufficient text for this to stand as a
validly-proposed taxon; ergo Buccinum maculosum Solander [Lightfoot,
1786] is a nomen nudum and unavailable for taxonomic nomenclature
(International Code for Zoological Nomenclature (ICZN) Article 12; Criteria of
Availability; see ref. below). How Dillwyn knew what Lightfoot had before him
is uncertain, but he may have seen the actual shell, which was to have been sold
at auction on Thursday May 25, 1786 according to the "Portland Catalog" (when
Dillwyn was in his ninth year of life) at some point. Needless to say, the
species is more than "very rare" in New Holland (Australia) as it does
not live there.
Dillwyn's other (second) citation was Seba vol. 3, pl. 68,
fig. 16 (see figure 2, taken from Müsch, I., R.
Willman, and J. Rust [eds]). Although the text of the non-binominal Seba work
is not available for taxonomic nomenclature, this illustration almost certainly
depicts the western Atlantic taxon in question - the editors' misidentification
of it as Tonna perdix (Linnaeus, 1758) notwithstanding. Dillwyn was
correct in recognizing his Buccinum maculosum as a new taxon!
Unfortunately, however, a quarter century earlier Gmelin (1791, p. 3476, no. 22)
applied that same binomen to a different shell, which we now know as the west
African Cassis tessellata (Gmelin, 1791) [Buccinum tessellatum
op. cit. p. 3476, no. 20]; see Abbott (1968; p. 54). The Dillwyn name is a
primary junior homonym of Buccinum maculosum Gmelin, 1791 and therefore,
until recently (see last paragraph),
unavailable for taxonomic nomenclature according to the ICZN Article 52;
Principle of Homonymy (ICZN, 1985).
Wait.... it gets more complicated! In looking further
into the synonymy in Tucker's Cassis monograph (Abbott, 1968; p. 54), I
saw that the binomen Buccinum maculosum was validly proposed by an even
earlier authority, Thomas Martyn (1784)! Because Martyn's Universal
Conchologist was not a consistently binominal work (like Seba), the
International Commission for Zoological Nomenclature exercised its plenary
powers (Anon., 1957) to declare most of Martyn's names invalid, while conserving
the names applied therein to nine well-known New Zealand marine gastropod species. View The Nine Species Among these
is Buccinum maculosum Martyn; now Cominella maculosa (Martyn,
1784). The consequence of this discovery is that B. maculosum Gmelin,
like B. maculosum Dillwyn, would not be not available (primary junior homonymy,
again; using ICZN, 1985). Since authors hadn't used B. maculosum Gmelin, quite possibly
because it was reported as a junior synonym of B. tessellatum Gmelin by
Dillwyn (1817; p. 596, no. 20), who appears to be the First Reviser (ICZN
Article 24). It is possible that Dillwyn chose B. t. over B. m.
because it appeared earlier in the Gmelin work (on the same page, two species
above it, a convention referred to as "position priority," employed by some
taxonomists in the past but never codified by ICZN) or perhaps because he
thought the earlier Solander [Lightfoot] (nude) name rendered Gmelin's
Buccinum maculosum unavailable. Whatever the machination, the outcome is
that the cassid (helmet shell), unlike our tonnid (tun shell), nomenclature
avoided being threatened with mass confusion by the loss of a such well-travelled binomen
as Tonna maculosa (Dillwyn), the topic of this discussion.
Wait; there's still more. Helix sulphurea C. B. Adams, 1849 (p. 33) was
first reported from Jamaica, and the author gave a short description, noted the
size (0.175 inch across by 0.16 inch high), and stated; "Perhaps this is a young
shell, but probably not of any described species." The next year he returned to
this taxon and wrote (C. B. Adams, 1850; page 98): "On page 33 we described
Helix sulphurea as 'perhaps a young shell.' This opinion was founded on its
papyraceous appearance. Mr. Chitty, from whom the original specimens were
received, has recently furnished a series, which shows that this shell, with all
the generic characters of Helix, is the last embryonic stage of the West
Indian variety of Dolium perdix!" [his exclamation point]. I believe
figure 3. is very much like what C. B. Adams had
before him in 1849 (and Chitty somewhat later). This is a photomicrograph of
the apex of the same shell as in figure 1. It shows an intact
ultraplanktotrophic protoconch reflecting a very long swimming veliger stage, an
attribute which no doubt accounts for the wide dispersal of this species and
many of its confamilial relatives. Acording to Clench and Turner (1950), the
type specimen is lost, but, given the quotes above, I don't thank anyone would
dispute the diagnosis of Helix sulphurea C. B. Adams, 1849! As far as I
can tell, the first reference to the synonymy of H. s. and Tonna
maculosa Dillwyn, 1817 [sic; no parens] is Turner (1948; p. 169).
It appears the next available name for the Atlantic
Partridge Tun is Dolium pennatum coined by the Danish conchologist Otto
Andreas Lowson Mørch (he actually signed this work "Mörch;" see below) from the
"Antill." It was published in another public auction catalog - that of the
Alphonso .... de Yoldi collection (its 4,012 lots rivaling the 4,156 of the
Dutchess of Portland). Mørch's indication (lot 2058; p. 110) was an
illustration in Martini - vol. 3, fig. 1078 (see figure 4).
This is interesting because Martini himself called it a juvenile of Buccinum
[now Tonna] perdix Linnaeus, 1758 and showed it alongside two
adult T. perdix! Nonetheless this juvenile shell shows the color pattern
of the W. Atlantic species (look at the smudgy pattern on the penultimate whorl
in figure 1) - not the Indo-Pacific T. perdix, with which it seems to
have been confused/associated by many authors. Ergo Tonna pennata (Mørch,
1852) refers to the Atlantic Partridge Tun, and the Partridge
Tun, T. perdix (Linnaeus, 1758) from the Indo-Pacific is closely related
- and not just biologically and by its vernacular name: Pennata probably
means "feathered" in reference to its color pattern, which a century before
likely inspired Linnaeus' perdix, Latin for "partridge."
Armed with all this
information about synonyms, homonyms, availability of nomina, etc., I was
getting ready to dismantle the underpinnings of the Atlantic Partridge Tun
nomenclature until a very perceptive correspondent of mine, Marien Faber
tactfully pointed out the value of "reading the directions" before launching
such an ambitious campaign. Marien indicated that the ICZN (1999;
fourth edition) contained an important policy change versus between the
third (1985) edition. Until 2000 a primary junior homonym (e.g. Buccinum
maculosum Gmelin, 1791; B. maculosum Dillwyn, 1817) was deemed
unavailable for taxonomic nomenclature (see end of
paragraph four) by the ICZN, and could only be conserved by petition to the
Commission to exercise its "plenary powers." The current (fourth; 1999) edition
states (Article 23.9.5): "When an author discovers that a species-group name in
use is a primary junior homonym of another species-group name also in use, but
the names apply to taxa not considered congeneric after 1899, the author must
not [our emphasis] automatically replace the junior homonym; the
case should be referred to the Commission for a ruling under the plenary power
and meanwhile prevailing usage of both names is to be maintained." Short
version: the burden of petition is shifted from authors who wish to conserve (a
majority of) primary junior homonyms to those authors who wish to suppress such
names. Ergo, until told otherwise, we are legit in using Tonna maculosa
(Dillwyn, 1817) as did the overwhelming majority of the authors who taught us
much of what we know of the Atlantic Partridge Tun.

Somewhat amazingly, along with another 40 other marine mollusks we've collected
in northeast Florida including the Lions-paw, this Tun is found in the eastern
Atlantic as well. See Vermeij and Rosenberg (1993) and Talavera (1982).
Lady Margaret Cavendish Bentinck, Duchess of Portland, died in 1785 leaving
extensive and valuable collections, not the least of which was of shells. The
Rev. John Lightfoot (1735-1788), her librarian and chaplain, compiled and
annotated a list of her curios. This 194 page, 4,156 lot inventory was printed
early the next year and served as the catalogue at the auction of her
collections conducted from late that May to June 18 (Lightfoot, 1786; Dance,
1966; Kay, 1965; Rehder, 1967). Allison Kay (1965; p. 10) gave convincing
evidence that Daniel Solander, a pupil of Carl Linnaeus (in 1761 ennobled as
Carl von Linné), a naturalist who curated material in the collection of the
Duchess, first penned many of the names employed by Lightfoot, but only in
manuscript form. Until 1965 most attributions for
the "Portland Catalogue" nomina were to Solander, but, after Kay's
report, the consensus among malacologists is that
Lightfoot is the proper authority. Although a substantial number of zoological names were
validly proposed by Lightfoot in this "Portland Catalogue," only a handful of
workers in the 18th and 19th centuries put them to use, often without proper
acknowledgement. Even the work of Sherborn (1902; later corrected), which
purported to list all the zoological names introduced from 1758 to 1800, listed
none of these names. Tom Iredale (1912) was the first to specifically wrest the names
from near oblivion. William Healy Dall (1921) investigated the "Portland
Catalogue" mollusks and presented selections of the Lightfoot text, which was
always brief, but often cited an illustration in an earlier work or two. A more exhaustive and definitive treatment was provided by Rehder (1967), who edited out 15 species names as nomina nuda (nude
names) due to lack of rigorous descriptions or cited illustration(s). The
author added five molluscan species names overlooked by Dall and proceeded to
pare the total (111) as follows: 39 junior synonyms (species validly named prior
to the "Catalogue"); three junior homonyms (names selected by Lightfoot but used
by prior authors); nine nomina dubia (descriptions and illustrations
insufficient to diagnose the species) and three nomina oblita (forgotten
names not employed in the literature between 1917 and 1967; this procedure has
become more involved since Rehder's work; see ICZN, 1985. Article 79(c)
Suppression of unused senior synonyms and ICZN,
1999. Article 81.2.2 and .3).
This reduction left 57 available
species and two generic names for Recent mollusks attributable to The Rev. John
Lightfoot. Rehder's systematic list at the end of his paper omitted the valid
species Subninella undulata and Strombus sinuatus, which were
given proper (and favorable) treatment in the text.
View An Emended List
Just to complicate matters a little more, Higo, Callomon, and Goto (1999;
G2833, p. 246) listed Buccinum
maculosum Gmelin, 1791 as a synonym of Colubraria muricata
(Lightfoot, 1786) [lot 2296, p. 104 of our now familiar "Portland Catalog"].
This is a lapsus calami for
Murex
maculosus Gmelin, 1791 (p. 3548, no. 79).
Lately revisionists (including Turgeon, Quinn, et al., 1998, who
also exalted Tonna pennata over T. maculosa) have worked hard to
render vernacular names in lower case. For a rebuttal see
http://www.jaxshells.org/flatcoil.htm.
Bibliography
Abbott, R. T. The helmet shells of the world (Cassidae). Part one.
Indo-Pacific Mollusca 2(9): 7-201. August 30.
Anon., 1957. Opinion 479. Opin. Int. Comm. Zool. Nomencl. 16: 365-456.
September 3.
Adams, C. B., 1849. Descriptions of supposed new Helicidae from Jamaica.
Contributions to Conchology 3: 33-38. October.
Adams, C. B., 1850. Descriptions of supposed new Helicidae from Jamaica.
Contributions to Conchology 6: 90-98. March.
Clench, W. J. and R. D. Turner, 1950. The western Atlantic marine mollusks
described by C. B. Adams. Occ. Pap. Moll. 1(15): 233-403 incl. pls.
29-49. June 26.
Dall, W.H. 1921. Species named in the Portland Catalogue. The Nautilus
34: 97-100; 123-132 (January; April).
Dance, S.P. 1966. Shell collecting, an illustrated history. Univ. of
California, Berkeley, pp. 1-34, 35pls.
Dillwyn, L. W., 1817. A descriptive catalogue of Recent shells. 2 vols.
J. and A. Arch, London. pp. 1-1092 plus index.
Gmelin, J.F. 1791. Systema naturae, ed.13, vol.1, part 6, pp. 3021-3909.
Higo, Callomon, and Goto, 1999. Catalogue and bibliography of the marine
shell-bearing mollusca of Japan Gastropoda - Bivalvia - Polyplacophora -
Scaphopoda. Elle Publications, Osaka. pp. 1-749.
ICZN (International Commission for Zoological Nomenclature), 1985.
International code of zoological nomenclature. Third edition. Univ. Calif.,
Berkeley, xx + pp. 1-338. Feb.
ICZN (International Commission for Zoological Nomenclature), 1999.
International code of zoological nomenclature. Fourth edition. I.C.Z.N.,
London. xxix + pp. 1-306.
Iredale, T. 1916. Solander as a conchologist. Proc. Mal. Soc. London 12(2,3):
85-93.
Kay, E.A. 1965. The Reverend John Lightfoot, Daniel Solander, and the Portland
Catalogue. The Nautilus 79(1): 10-19. July.
[Lightfoot, J.], 1786. A catalogue of the Portland Museum, lately the
property of the Dutchess Dowager of Portland, deceased, which will be sold at
auction.... London, viii + pp. 1-194.
Linnaeus, C., 1758. Systema naturae per regna tria naturae... editio decima,
reformata. Vol 1. Regnum animale tenth edition. Stockholm. i+ pp. 1-823.
Martini, F.W., 1777. Neues systematisches Conchylien-Cabinet... vol. 3
pls. 66-121. Nuremburg. [All plates with annotations reprinted in Richardson,
L., R. T. Abbott, G. M. Davis [eds.], Early references to the figures in the
Conchylien Cabinet of Martini and Chemnitz: volumes I - XII. Tryonia
2, parts 1, 2: pp. 1-427, 432 pls., 70 vigns., August 1, 1979.].
Martyn, T., 1784-87. The universal conchologist....2 (some 4) vols.
London, 27pp., 80pls. [not seen].
Mörch, O. A. L. M., 1852. Catalogus conchyliorum quae reliquit D. Alphonso
D'Aguirra & Gadea Comes de Yoldi, regis Daniae cubiculorum princeps, ordinis
Dannebrogici in prima classe & ordinis Caroli Tertii Eques vol. 1 Cephalophora.
Hafnia (Copenhagen): pp. 1-170. August.
Müsch, I., R. Willman, and J. Rust [eds], 2001. [Seba, A., 1734-1765] Cabinet
of Natural Curiosities Locupletissi rerum naturalium thesauri. Taschen,
Köln. pp. 1-588 incl. numerous plates.
Rehder, H.A. 1967. Valid zoological names of the Portland Catalogue. Proc.
U.S.N.M. 121(3579): 1-51.
Sherborn, C.D. 1902. Index animalium sive index nominum quae ab AD MDCCLVIII
generibus et speciebus animalium imposita sunt. London. [not seen].
Talavera, F. G., 1982. Los moluscos gasteròpodos amfiatlànticos (estùdio
paleo y biogeogràfico de las especies bentonicas litorales). Colecciòn
Monografias 10 , Secretariado de Publicaciones de la Universidad de La Laguna,
La Laguna, Canary Is., pp. 1-351.
Turner, R. D., 1948. The genus Tonna in the
western Atlantic. Johnsonia 2(26): 165-192. October 30.
Turgeon, D. D., J. F. Quinn, Jr., A. E. Bogan, E. V. Coan, F. G. Hochberg, W. G.
Lyons, P. M. Mikkelsen, R. J. Neves, C. F. E. Roper, G. Rosenberg, B. Roth, A.
Scheltema, F. G. Thompson, M. Vecchione, and J. D. Williams, 1998.
Common and scientific names of aquatic invertebrates from the United States and
Canada: mollusks, 2nd edition. American Fisheries Society, Special
Publication 26, Bethesda, Maryland.
Vermeij, G. J. and G. Rosenberg, 1993. Giving and receiving: the tropical
Atlantic as donor and recipient region for invading species. Amer. Malac.
Bull. 10(2): 181-194. Dec. |