Color-full and eye-catching: an iconography of Liguus land ... · Polymita. are colorful and plentiful — at least in those days —, and were desired objects for conchologists

eschweizerbartxxx sng-

Arch. Molluskenkunde | 143 | (1) | 1–19 | 9figures | FrankfurtamMain,27.06.2014

© E. Schweizerbart’sche Verlagsbuchhandlung (Nägele u. Obermiller), 2014, ISSN 1869–0963DOI 10.1127/arch.moll/1869-0963/143/001-019

Authors’ adresses:1 Abraham S.H. Breure Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, the Netherlands; E-mail: [email protected]; *corresponding author2 Luis Álvarez-lajonchère, Museo de Historia Natural ‘Felipe Poey’, Facultad de Biología, Universidad de La Habana, El Ve-dado, Plaza, La Habana, Cuba; E-mail: [email protected] Adrián GonzÁlez Guillén, Coral Gables, U.S.A; E-mail: [email protected]

Color-full and eye-catching: an iconography of Liguus land shells

(Gastropoda: Orthalicidae)

aBraham S.H. Breure1 *, luis Álvarez-lajonchère2 & adriÁn GonzÁlez Guillén3

Abstract

The first book illustrating a Neotropical land shell appeared in 1684, with a figure of Liguus virgineus (linnaeus 1758). An iconography for this genus is presented, describing early pre- and post-Linnean figures from 17th and 18th century shell books, with a briefer discussion of later il-lustrations through the early twentieth century. Also the development until the first photographs in the early 20th century is briefly discussed.

Key words: history of malacology; pre-Linnean illustrations; Neotropical land shell; Orthali-cidae.

Introduction

Shell collecting has a long history dating back to pre-historic times. The earliest writings on mollusks were those about marine species and are attributed to Aristotle (dance 1966), while the first book figuring a freshwa-ter mussel dates back to 1491 (valledor de lozoya & araujo 2011). Land snails have attracted less atten-tion and Neotropical species became only well-known after trade connections had been established with e.g. the West Indies. Specimens from genera as Liguus and Polymita are colorful and plentiful — at least in those days —, and were desired objects for conchologists.

The figures of shells in natural history books have been the basis of identification for individuals who did not have access to a museum collection where the spe-cies concerned was represented. The quality of the figure was thus important to reflect all the characteristics of the species. In the case of Liguus, e.g., the overall shell shape, the shape of the columella and the colors were among some of those characteristics. However, the im-age quality can only be appreciated with some knowl-

edge of the printing techniques used at the time the book was produced (Buchanan 1979). Before 1700, the wood-cut was the earliest method which produced images in black and white. The image had to be drawn and carved in the wood block in reverse, so that when the inked block was pressed to a sheet of paper the image would be displayed correctly. When printers (or engravers with later techniques) forgot to do this, the shell appeared as mirror-image. In the course of the 17th century wood en-graving was developed as a refinement of wood-cutting that allowed to obtain a multitude of fine lines and thus subtle gradations of grey tones. Engraving a figure de novo was a laborious work, and it was thus not uncom-mon to re-engrave already published illustrations (Ford 2003).

Around 1700, copper engraving was introduced, which enabled illustrations colored by hand; this method was used throughout the 18th and into the early 19th century. The original drawing was transferred to a cop-per plate in reverse, after which the lines were cut away


2 Breure, A.S.H. et al.: Iconography of Liguus land shells

with a burin, and the plate was inked and printed. In this and the preceding technique the engraver worked from light to dark; a reverse technique (mezzotint) was developed in Germany in the 17th century in which the surface was roughened before the drawing was trans-ferred in reverse with a scraper. This surface technique was better suited for color printing than line processes, but the drawback was that the plates wore each time a print was taken from them and hence no two prints were exactly the same. Often the plates needed retouching, making further differences between the prints, until the whole plate had to be re-modeled.

Another technique made use of etching rather than engraving, in which the plate was covered with var-nish first, then etching needles and punches were used to copy the design with small dots which went right through to the copper beneath. Subsequently acid was applied eating away through the copper in which the dots had been punched, leaving intact only the parts covered with the varnish. This way, a very delicate and varied tone was produced, which was generally used with color printing on the plate.

In the aquatint technique the main difference was that the etching ground was porous. Acid was applied to the plate many times and this method produced a soft half-tone effect. As with mezzotints, the plates wore and had to be continually worked on, and many prints contained a number of minor differences. During the Grand Era of natural history books (1780–1830) copper engravings

were still used, but the more important works were il-lustrated with stipple engravings in France, and with aq-uatint or mezzotint in Britain. Color printing from plates was done either by applying all the colors to a single copper plate (Britain) or making several identical copper plates and applying different colors to each (France).

After 1830 these techniques were gradually replaced by lithography, and finally chromolithography. Lithographs used a smooth stone on which the drawing was made in pen and greasy ink or a greasy lithographic crayon. The soft-ness of the crayon gave a rather imprecise line, frequently improved by retouching. The stone was then wetted, but the inked parts rejected the moisture; when a greasy ink was passed over the stone it was, however, accepted by the previously inked portions. The lithographic prints, contrary to the aforementioned techniques, did not have the plate mark surrounding the engravings and were frequently hand-colored. In chromolithography one used as many stones as there were colors to be applied, which allowed for overlap-ping blocks of color overlapping and overprinting to create a range of colors. However, this process required a con-siderable skill of the printer and many chromolithographic plates were not of the highest quality (Buchanan 1979: 193–197). The introduction of photography superseded these techniques in the early 20th century.

In this paper we will describe how the Liguus shells were illustrated drawn in pre-Linnean times and beyond, until in the early 20th century when the first photographs appeared.

Iconography of Liguus shells

The first published figure in print of a Liguus shell we have found is are aware of was in PhiliPPo Buonanni’s [also known as Filippo Bonanni] Recreatio mentis et oculi (Buonanni 1684). This is the earliest book devoted entirely to shells (dance 1966), and a typical late 17th century compendium. The folio volume, which invited contemplation rather than reading, contained copper-plate engravings illustrating individual shells. Each il-lustration was accompanied by a relatively brief text which described the shell in terms of its external fea-tures, enabling the classification of objects in conchol-ogy (dietz 2006). In Classis III, Testacea Turbinata, figure 66, Buonanni pictures a dorsal view of a shell that is now attributed to L. virgineus (linnaeus 1758) (PilsBry 1899), although linnaeus did not mention this figure until the 12th edition of his Systema naturae (lin-naeus 1767 [1766-1767]). The shell is placed in a semi-upright position and —though reversed —shows clearly a Liguus species (Fig. 1A). This figure has been copied in Buonanni (1709).

The explanatory text reads (Buonanni 1684: 121): “Quinque fascijs cingitur hic alius caeteris elegantior.

Altera colore subnigro est, altera purpurea, vel ad min-ium accedens, tercia fulua [fulva], viridis quarta, vltima ostrina; omnes ita à summo vertice mucronem vsque ita circumuoluuntur, vt interuallo [intervallo] quodam distinctae inter se, in eo videatur veluti album indusium, quò turbo tegitur. Valdè nitidus est, & lenigatus, & ab Indico mati cum caeteris octo superioribus habetur” (A liberal translation might be: “Encircled with five bands and otherwise quite elegant. Another color is semi-black [grey], or purple, or tending to vermillion, as a third brown, as a fourth green, real purple as the last one, all wrapped up to and including the apex, with clear inter-ruptions, which seems like it were a white coat covering the whorls. It is very shinying, [...], and on the testimony of the sea with the eight other [species] above”). The shell shape in the figure alone is, however, distinctive enough: the only Liguus sp. that has a high, near 40º angle is L. virgineus. Pl. 151 fig. 1 in Petiver (1764) is a slightly changed, mirror-copy of Buonanni’s figure.

Shortly after this publication, martin lister’s Histo-riae sive Synopsis Methodicae Conchyliorum appeared (lister 1692–1697; see also Wilkins 1957); a book not


Breure, A.S.H. et al.: Iconography of Liguus land shells 3

Figure 1. A—Buonanni 1684: Cl. III, fig. 66; B— lister 1692–1697: pl. 12 fig. 7; C— lister 1692–1697: pl. 15 fig. 10; D—klein 1753: pl. 2 fig. 43; E—arGenville 1742: fig. M–N; F— arGenville 1757: App., pl. 1 fig. G; G—müller 1766: fig. 52; H—schröter 1779: pl. 8 figs 3–4.



Figure 2. A—Gualtieri 1742: I, pl. 6A; B— Gualtieri 1742: I, pl. 6D; C— Gualtieri 1742: I, pl. 6C; D— arGenville 1780: pl. 65.

only figuring shells, but also animals and anatomical details. The (as such unnumbered) plates in this work are composed of illustrations that have been partly grouped together, to show multiple shells on a page. As a result there is a dual numbering, one close to the shell (here indicated as “figure”, and repeated with the explanatory text), and one to the lower-right of the shell (here used as indication for “plate”). Plate 15 fig. 10 (Fig. 1C) shows a Liguus shell, for which the text reads “Buccinum septem orbuun, fascys fere bicoloribus interdum tricoloribus undis instar depictum” (Liberal translation: Whelk of seven whorls, the waving lines are bi- or tricolored). The figure shows the shell laying flat on a background from a lateral-posterior angle, which does not show the shape of the aperture very well. However, it shows the widely concave anterior lip, or `base´of the shell, in a way that one can easily see the columella if looking from bellow, which is exclusive for L. virgineus. It may be noted that in lister’s classification the color pattern of the shells played an important role (cf. dietz 2006: 367). This is

the shell referred to by linnaeus (1758: 740) under his Buccinum virgineũ. lister’s (1692–1697) Plate 844 fig. 72 —cited by Gmelin (1791: 3429) —is very dubiously referred to this species. There is a second figure in plate 12 fig. 7 (Fig. 1B), as “Buccinum septem spirarum, cujus superiores spirae fasciatae inferiores undatae” (“Whelk of seven whorls, upper ones with bands, the inferiors with waves”); note that here the shell is described with the apex pointing down, as they were sometimes illus-trated in those days, see below). The figure shows the shell from a lateral side, with the plane of the aperture slightly skewed and turned towards the viewer. müller (1774: 146) quoted this figure, among others, for his third variety of B. fasciatum: γ alba fascies variegatis (white, lines variegated). This shell is a variety of Liguus fasciatus (müller 1774), viz. L. f. goodrichi clench 1934. The same two figures were reproduced in the Hud-desford edition (lister 1770); the first figure was copied by Petiver (1702–1704) in his Gazophylacii naturae et artis, pl. 22 fig. 11 and re-issued in Petiver (1764).



Figure 3. A—knorr 1757: pl. 30 fig. 7; B—reGenFuss 1758: pl. 10 fig. 46; C—chemnitz 1786: pl. 117 fig. 1000; D— chemnitz 1786: pl. 117 fig. 1002–3; E— chemnitz 1786: pl. 117 fig. 1001; F— seBa 1759: pl. 40 fig. 38; G—seBa 1759: pl. 40 fig. 38[bis].

nicolò Gualtieri, in the first part of his Index Testarum Conchyliorum (Gualtieri 1742), figured a number of taxa, which he regarded as “Buccinum fluviatile”. These spe-cies are figured as dorsal and ventral views of the shell placed on its apex, their aperture plane perpendicular to the viewer. This upside-down illustration of shells was not uncommon in those days, particularly in France (Ford

2003). For Gualtieri the shape and position of the aper-ture was part of his classification system, hence the illus-tration of the aperture in the position as we use them till today. Among these shells are those of three Liguus taxa (Gualtieri 1742: I, plate 6 figs A, C and D; Figs 2A–C).

The explanatory text of his figure A reads: “Bucci-num fluviatile, spiris non prominentibus oblongum, ore



Figure 4. A— seBa 1759: pl. 39 fig. 73; B— seBa 1759: pl. 39 fig. 74; C— knorr 1757: pl. 25 fig. 4; D— chemnitz 1786: pl. 117 fig. 1004; E— chemnitz 1786: pl. 117 fig. 1005; F— chemnitz 1786: pl. 117 fig. 1006.

angustiore, laeve, candidum, fasciis aliquando piceis, aliquando rubris, plumbeis, & luteis, iridis instar elegan-tissime fasciatum” (Freshwater whelk, oblong with in-conspicuous whorls, with a narrow aperture, thin, white, sometimes has black bands, sometimes red, livid or yel-low, with evenly spaced and tasteful bands). The misin-terpretation of Liguus species as freshwater shells was evidently due to the incomplete data with which the European conchologists received their material from overseas. This shell has been considered as L. virgineus, and the figure was referred to by linnaeus (1758: 740). However, sWainson (1822 [1821–1823]: pl. 84 middle figures + text), referring to the same figure of Gualtieri, considered it to be his Achatina vittata [= L. vittatus (sWainson 1822)]. We concur with this, as the narrow aperture to which Gualtieri refers hints more to L. vit-tatus than to L. virgineus, which has the aperture with a

wide, almost flat basal line. Although Gualtieri’s figure clearly illustrated L. vittatus, the description seems to mix the two taxa when describing the color bands; livid and yellow are typically colors of L. virgineus.

Plate 6 figs C–D (Gualtieri 1742) shows two subspe-cies of L. fasciatus. The text of figure C is: “Buccinum fluviatile, maius, laeve, labio interno repando, ex carneo, fulvo, albido, & purpurascente colore fasciatum, ali-quando lineis intersectis, punctatum, nebulatum, & mar-moris intar lucide, & eleganter variegatum” (Freshwater whelk, bigger, thin, with the inner lip curved, sickle-shaped colored in flesh [reddish] color, brown, white and purplish, sometimes with broken lines, punctured, smudged, bright or marbled on the inside and tastefully colored). Given the apparent size, eight whorls, long spire, the concave and very truncated columella, this is a shell of the L. fasciatus achatinus lineage, a dark morph,



Figure 5. seBa 1759: pl. 39. Reproduced with permission from the copy in the Koninklijke Bibliotheek, The Hague.



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probably found near Trinidad, Cuba. Gualtieri’s figure was referred to by linnaeus (1767 [1766–1767]: 1186) as “[var.] β” of B. virginea. This is the first illustration cited for Buccinum fasciatum var. γ (müller 1774). Fi-nally, the explanation of figure D reads: “Buccinum flu-viatile, idem minus, candidum, & in prima, & in secunda spira linea subrubra circumdatum” (Freshwater whelk, as small, white, and first and second whorl with a red-dish spiral band). This is L. fasciatus pallidus (sWainson 1822), and was cited by müller (1774: 146) for his vari-ety β of Buccinum fasciatum.

In the third part of Gualtieri’s work, in the section on Buccinum, plate 45 fig. D illustrates “Buccinum parvum, integrum, ore obliquo, umbilicatum, laeve, albidum, duabus lineis rufis circumdatum” (liberally translated as: Small whelk, with an oblique aperture, umbilicate, smooth, whitish with two spiral red lines). This figure is cited as Liguus virgineus by both müller (1774: 143) and Gmelin (1791: 3429).

antoine josePh dezallier d’arGenville published his L’Histoire naturelle (also known as Conchyli-ologie) in the same year (1742) as Gualtieri’s book. d’arGenville’s work has been both praised and criti-cized. On one hand, “were it not for his illustrations, his further comments would be quite unintelligible” and “criticized severely in his lifetime” (dance 1966: 59). On the other hand, “this illustrated compendium of shells enjoyed an extraordinary success” and was “the founda-tion on which the magnificent Paris collections of the eighteenth century rested” (dietz 2006: 368). On plate 14, under the heading “Vis” and amidst a variety of ma-rine species, two figures of Liguus may be found (figs M–N; Fig. 1E); both figures are with the apex up. One is a finely engraved, dorsal view of a Liguus fasciatus vari-ety (arGenville 1742: pl. 14 fig. M). The accompanying text (arGenville 1742: 276) reads: “on sera seulement remarquer que celle de la lettre M, apellée le Ruban, est bariolée seulement par le haut, & toute blanche par le bas, à la difference des Vis ordinaires, que l’on apelle Rubans, comme celle marquée N”. Again at least sev-en whorls, a globose, white last one, a flatter profile of the others, the only brightly colored (bariolée), and the trimmed bands just intensified at the ends point towards a L. fasciatus achatinus like shell. müller (1774) com-bined this L. f. achatinus, and the L. f. pallidus of Gual-tieri pl. 6 fig D, in his β variety of Buccinum fasciatum; while he paid perhaps too much attention to the paler last whorl, in this case he was wrong.

The second figure [arGenville 1742: pl. 14 fig. N] shows a somewhat smaller shell. The plane of the ap-erture is here skewed, turned away from the viewer, thus appearing very narrow and rather elongate. lin-naeus (1758: 740) referred to this figure for his Bucci-num virgineum; also mentioned in linnaeus 1764 (: 612) and 1767 (1766–1767: 1186). d’arGenville (1742: 382) stated “les Vis apellées rubans, venant des Barbabes”; this locality refers back to lister (1692–1697). This is

indeed a specimen of L. virgineus, not only for the posi-tion of the three lines, or the name “ruban” [= banded as the locals use to call it], but because of the excavated concavity of the base of the columella clearly shown, which is typical of this species.

In the second edition (arGenville 1757) a newly en-graved figure appears in the Appendix, pl. 1 fig. G (Fig. 1F), which is a re-design of arGenville 1742: fig. M; it thus represents a free interpretation of L. fasciatus. The accompanying text (arGenville 1757: 384) described the shell as “Huit bandes principales, assez larges, con-vrent cette Vis: la première est brune, la seconde ver-dâtre, la troisiéme violette, la quatriéme bleue, la cin-quiéme blanche, bariolée de brun rouge, la sixiéme à peu près de même; la septiéme & la huitiéme formant le sommet, sont de couleur de rose”. This shell, from d’arGenville’s own collection, is Liguus virgineus.

jacoB theodor klein (1753) wrote an extensive clas-sification of shells, today almost forgotten, in which he recognized a genus Oxystrombus, with 11 species ar-ranged in two groups, making references to figures of Buonanni and lister. One of his species in the group Laevis was described as “Fasciatus supra; infra septem spiris undatus; ore subrotundo laevi”; his pl. 2 fig. 43 (Fig. 1D) is a modified copy of lister’s Liguus fasciatus. This figure is the last quoted under müller’s Buccinum fasciatum var. γ: alba, fascies variegatis müller (1774: 146). klein’s taxon Oxystrombus was referred to by mörch in his Catalogus Conchyliorum (post-Linnean and validated for nomenclatural purposes; mörch 1852: 21) for the species group of Liguus fasciata (müller), L. crenatus (sWainson 1821), and a “var.[iety] l[on]g. 73 m[illimeter] A.[chatina] fasciata var. ع Pf[eiffe]r [in] reeve [reeve 1849 (1849–1850): pl. 10] fig. 35 c”, which is L. f. achatinus. mörch (1852) kept this group apart from Pseudotrochus ‘klein’ herrmannsen 1847 = Chersina ‘humPhrey’ Beck 1837 = Liguus montFort 1810, in which he only listed L. virgineus; this was based on klein’s figure of this taxon, listed as “PseudoTro-chus” (klein 1753: pl. 7 fig. 116). Since mörch included only subspecies of L. fasciatus and no other species un-der Oxystrombus, he thus correctly differentiated the two taxa (groups of species) and this could be understood intending ‘subgenera’, because Oxystrombus is printed in a smaller, thinner type and between parenthesis, while Liguus, which is printed in bold type, is listed above.

The first colored figures appeared in GeorG WolF-GanG knorr’s Vergnügen der Augen und des Gemüths. This book consisted of several parts, originally published 1757–1771, and re-published in a French version (knorr 1760–1773) and a Dutch version (knorr 1770–1775). Each part contained merely a description of the many plates that showed shells in no particular order. In part 1 — published 1757 — a dorsal view is shown of a shell on pl. 30 fig. 7, in which Liguus virgineus may be recognized (Fig. 3A); the figure is not very precise, but the explanatory text leaves no doubt about the identifi-



Figure 7. seBa 1759: pl. 39 fig. 62–72.

cation. The same figure was re-published in the French (1760) and Dutch (1770) versions of the book. In part 5 pl. 25 fig. 4 — published in 1771 and re-published in the French (1771) and Dutch (1774) versions —a second Liguus shell is figured (Fig. 4C). It is a dorsal view, with the axis of the shell slightly turned to the left, of what nowadays would be classified as L. fasciatus goodrichi clench 1934: only brown and white with the subperipheral and subsutural lines reddish-brown and

rose apex (the rose columella not shown). This accurate illustration is not referred to by müller, nor sWainson. A third shell was published in the sixt part (also 1771), pl. 28 fig. 4; Gmelin (1791: 3429) referred to this figure as Liguus virgineus. The latter two shells were ‘Ex Museo Houttuijniano’, thus from the collection of martinus houttuyn, who contributed much to the parts published after knorr’s death in 1761. See Boeseman & de liGny (2004: 68–71) for details on houttuyn’s contributions.



Figure 8. PilsBry 1912: pl. 37.



Franz michael reGenFuss, in a bi-lingual German / French edition, published in 1758 a Liguus species, which he referred to as “Buccinum terrestre f. fluviatile”. Besides references to the previous literature, for the first time we find here vernarcular names in three languages (Danish, German and French). His pl. 10 fig. 46 (Fig. 3B) pictures two views of what seems to be Liguus f. fasciatus from the Havana-Matanzas region; the rounded profile of the whorls, seven (not eighth) turns, shorter spire, the bigger last whorl with its globose shoulder, not at the base, the arched but barely twisted or truncated columella, the mauve parietal, and the blue-grey tint on the upper whorls, are all characteristics more frequently found on these shells. The left hand picture is a shell in ventral view, but with the axis turned to the right-hand side at 45 degrees. The colors are vivid and the shell is shown to be very shining. The right hand picture shows a dorsal view, the shell with the aperture placed on the underground and the axis to the left-hand side at 315 degrees. See van Benthem juttinG (1964) and dance (1966: 59–60) for the history and importance of this work within its epoch.

The third book with colored figures of Liguus was published by alBertus seBa in his Locupletissimi re-rum naturalium thesauri accurata descriptio. This work started in 1734 and was issued in four parts, each in two bi-lingual editions (Latin / Dutch and Latin / French). Part of the editions was issued with hand-colored plates (see below). The third volume, in which Liguus appears, is generally cited as seBa (1758). However, holthuis (1969) has shown that this posthumous volume, edited by vosmaer, was published — after an eventful his-tory — in 1759. The correct citation is thus seBa (1759). seBa’s work is renowned for its artful illustrations, which reflected the arrangements in his collection (Figs 5–6). linnaeus, in his various editions of the Systema Naturae, referred many times to these ‘pre-Linnean’ illustrations, some of which have been cited as type by later authors (e.g., Bauer 2003, crochet et al. 2006, kohn 1963; see also Boss 1988, heller 2007, and below).

seBa’s Plate 39 shows an arrangement of various shells, more or less geometrically positioned, on which figures 62–74 represent the idiotypes of Liguus fasciatus (Fig. 4A–B, 7), quoted by müller (1774: 145). Figs 62–68[a], 70–71, 73 are dorsal views; figs 68[b]–69, 72 and 74 are ventral views. Figs. 62 and 63 are the first idiotypes for var. α of Buccinum fasciatum, fig. 67 is the last for his var. β, and fig. 68 for the γ; being all color morphs of the same subspecies, this was incorrect. Various figures are probably illustrating the same shell in ventral and dorsal views: figs. 68[a–b], 69–70, 71–72, 73–74; figs. 58–59 have not been referred to by later au-thors but represent an albino L. fasciatus shell. The fig-ures on the right-hand side of the plate are mirror-images of the shells, as the coiling of the shells in the right-hand figures is wrong; this is likely due to an error in the en-graving of the copper plate template as enantiomorphy is

this species is very rare. PilsBry (1929) selected figure 74 as type figure of L. fasciatus (Fig. 4B); this is one of the mirror-images. All of them belong in what we known today as L. f. guitarti jaume 1952, a Sancti Spiritus mi-crogeographical race of L. f. goodrichi from southern Cienfuegos, Cuba. On pl. 40 fig. 38[a–b] represents the second known Liguus species at that time, L. virgineus. The left-hand figure gives a dorsal view, the right hand figure a (mirror-imaged) ventral view (Fig. 3F).

In 2001, the plates from the hand-colored copy of seBa’s work, present in the Royal Library in The Hague, were reproduced with modern printing techniques (müsch et al. 2001). As Bauer (2002) correctly stated, the warm colors of the original pigments and the depth of the copperplate engravings cannot effectively be cap-tured by these techniques. müsch et al. (2001: 23) as-sumed that the volumes were not sold in colored condi-tion, but rather that individual buyers commissioned colorists independently to color the plates. The plates of the third volume, except pls 32–34, were already made during seBa’s lifetime (holthuis 1969: 241–242). There-fore, it is likely that the colorists based themselves on previously colored copies if they were ordered after the publication of this third volume in 1759. This is cor-roborated by the two copies used for the 2001 reprint, as there “are striking differences in the way that these two copies have been painted” (müsch et al. 2001: 544). se-Ba’s collection itself was auctioned in 1752, and widely dispersed afterwards (Boeseman 1970).

linnaeus had visited seBa in 1735, when the second volume of the Thesaurus was published, and used many of seBa’s animals as type specimens for his own descrip-tions (Eason 2014). linnaeus (1758: 740), in describing “Buccinum” virgineus, cited six (6) illustrations, i.e., lis-ter (pl. 15, fig. 10); Petiver (pl. 22, fig. 11); Gualtieri (pl. 6, fig. A [= Liguus vittatus]); d’arGenville (pl. 14, fig. N); klein (pl. 7, fig. 116), and reGenFuss (pl. 10, fig. 40 [sic, 46]). In the 12th edition a reference to seBa’s fig-ures and a variety β [= L. fasciatus] was added. linnaeus erroneously recorded this species from “Africae fluviis” (African rivers). See Table 1 for a full cross-reference of linnaeus’ citations to present-day taxa.

In 1766 Friedrich heinrich Wilhelm martini pub-lished a paper, in different parts, on land snails in the Berlinisches Magazin, oder gesammelte Schriften und Nachrichten für die Liebhaber der Arzneywissenschaft, Naturgeschichte und der angenehmen Wissenschaften überhaupt. In the third part (martini 1766: 125) he refers to d’arGenville’s earlier work (arGenville 1757). The shell figured in his pl. 5 fig. 52 (Fig. 1G) is a copy of d’arGenville’s pl. 1 fig. G, and thus represents Liguus fasciatus (but very badly). This paper is of interest since it is the first in which a comparative discussion on Liguus literature was made from a scientific point of view. How-ever, martini mistakenly attributed the figured specimen to Jamaica, relying on data of the collector, where this genus does not occur.





Figure 9. simPson 1920: frontispiece.

An obscure reference was found in chemnitz (“En-cyclop. Rec. de Pl. tom. 6 tab 64 f. 2”, which in the end could be traced back to Anonymous (1768). The expla-nation of pl. 64 fig. 2 reads “Celui de la fig. 2. se nomme le Ruban, parce qu’il a des bandes circulaires blanches, jaunes, vertes & rouges qui imitent assez bien un ruban: on le trouve à Saint-Domingue [Hispaniola]”. The figure is very poor, but the text undoubtedly refers to Liguus virgineus.

schröter, in his Geschichte der Flussconchylien (1779), published a lengthy discussion of the literature on Liguus virgineus, still considering it a freshwater snail. He noted the considerable variation in color pat-tern of the shells then known, and discussed the two different vernarcular names — “Staatenflagge” and “Prinzenfahne” — that were in use (schröter 1779: 335–339). In plate 8 figs 3–4 two figures are presented, both up-side down and in monocolor, of dorsal views of this species (Fig. 1H).

In 1780 jacques Favanne de montcervelle and his son published an enlarged edition of d’arGenville’s Conchyliologie (arGenville 1780), in which the shells were figured as arranged in an artful manner in cabinets. On plate 65 six Liguus shells are arranged in a star-like fashion, their apices pointing together (Fig. 2D). Figs G1 and G4 present a ventral view of Liguus fasciatus var. crenatus (sWainson 1821), one is a dextral shell (G1), the other a sinistral one (G4). However, since sinistral shells are very rare in natural populations, it is likely that the design was made with artistic rather than scien-tific purposes; therefore, we assume that G4 is a mirror-image. The other four shells are figured as dorsal view. Two of them (G2 and G6) represent perhaps L. fasciatus fasciatus, the others (G3 and G5) look more like L. f. achatinus, but the whole figure is rather imprecise.

martini started in 1769 with his Neues systematisch-es Conchylien-Cabinet, a work in which he intended to illustrate every shell known, but in a systematically or-dered way. After his death in 1778, johann hieronymus chemnitz continued the work which, with various edi-tors and authors, would last till the early 20th century. In 1786 chemnitz published a volume on land and freshwa-ter mollusks, with contemporary rather good illustrations and very brief explanatory text. On plate 117 (chemnitz 1786) we find the two Liguus species known at that time; the text uses the binominal system and gives the Ger-man vernarcular name. Figures 1000–1003 “Die Staaten-flagge. Die Prinzenfahne. Bulla virginea Linnaei” (Fig. 3C–D). Figures 1004–1006 “Der vielfarbichte Bund. Bulla fasciata” (Fig. 4D–F). The latter depict either L. f. guitarti jaume 1952 or L. f. poncianus sanchez roiG 1951.

In the tenth volume (chemnitz 1788), on pl. 173 figs 1682–1683, Liguus virgineus was illustrated again; these figures have been cited by Gmelin (1791: 3429) as vari-ety δ.

In 1791, Gmelin published the 13th edition of the Systema Naturae, and under “Bulla” virginea (linnaeus 1758), Gmelin listed (1791: 3429) four of the six sources cited by linnaeus, along with “Spengl. selt. Conch. t. 1 fig. G” [?arGenville 1757]; schröter ([1779] pl. 8, fig. 3–4), schröter ([1783] pl. 5, fig. 7), knorr (6, pl. 28, fig. 4), Favanne ([arGenville 1780] pl. 65, fig. G); and chemnitz (9, pl. 117, figs. 1000-1003). Gmelin also recognized four varieties, each named as polynomials (invalid), and each based on one or more illustrations, including Buonanni (cl. 3, fig. 66). Gmelin’s locality for this species was even more incorrect than the locality given by linnaeus, recording it from “Asiae fluviis” [a freshwater species from Asia].

Gmelin (1791: 3430) also listed “Bulla” fasciata mül-ler, and cited five illustrations for that species; as local-ity he cited “America australis et India”. He recognized five varieties, each named as polynomials (invalid), and each based on one or more illustrations. The latter two are herein excluded from the list of references to this species. See Table 1 for a summary of all references up till this publication.

The work of montFort (1810: 422) defines Liguus just for L. virgineus by monotypy; the figure is very poor, with too many more whorls, the wrong profile, but the base of the columella is well depicted. The descrip-tion of the species makes it likely that he had the correct shell at hand, although all localities cited were wrong.

During the 19th century the engraving and printing techniques improved to such an extent, that iconographic works with extensive systematic treatments of species became fashionable; e.g., sWainson’s Zoological illustra-tions (sWainson 1821–1823), the Conchologica Iconica (reeve 1848–1850, 1849–1850), and martini und chem-nitz’s Neue Systematischen Conchylien-Cabinet, Neue Folge, which started in 1837 and lasted till 1920 (küster & PFeiFFer 1840–1865; see coan et al. 2012a, b for a full collation of these references).

sWainson’s work is paramount in the early 19th centu-ry. His plate 42 depicts one of his new taxa, viz. Achati-na pallida sWainson 1821 (= L. f. var. pallidus) in lateral views of both the dorsal and ventral sides; pl. 58 another one, viz. A. crenata sWainson 1821. Plate 74 illustrated A. fasciata sensu müller (currently understood as the variety L. f. guitarti), both in a skewed dorsal and more or less ventral view; pl. 84 showed A. vittata sWainson 1821 and A. emarginata sWainson 1821; the latter is ac-tually a variety of L. virgineus. Finally, his pl. 162 show-



ing L. f. goodrichi, L. f. achatinus (ventral and dorsal views; both are from the eastern provinces in Cuba), and a juvenile of L. f. fasciatus or maybe L. f. pallidus.

Another work within this era relevant to understand the history of Liguus illustration, is the publication by orBiGny (1841–1853). Plate 6 shows the following forms: Figs 1–2 is L. f. trinidadense jaume 1952 in ven-tral and dorsal view; this shell has very large proportions and a pink columella. Figs 3–5 is according to orBiGny the typical Achatina fasciata of müller; in our cur-rent understanding fig. 3 is L. f. guitarti jaume, fig. 4 L. f. poncianus sanchez roiG, and fig. 5 L. f. goodrichi clench. Fig. 6 is L. f. pictus reeve 1849, probably a morph from Key West in Florida (if displayed ventrally the shell should have a pink columella). Fig. 7 is a vari-ety of L. f. solidus say 1825 like humesi jones 1979, or versicolor simPson 1920 from Florida. Finally, fig. 8 is the morph L. f. crenatus sWainson from, given the size, the eastern region in Cuba.

reeve’s work is interesting as his plates were hand-colored, but the quality of coloring varies considerably from one copy to another (Petit 2007: 27). On plates 7, 9 and 10 of his treatise on Achatina (reeve 1849 [1849–1850]) several forms of Liguus may be recognized. Plate 7 figs. 22 a–b are currently considered as what the leg-end reads, Liguus murreus reeve 1849. clench (1932) wrongly associated reeve’s fig. 22b with L. f. solidus (say 1820) var. roseatus PilsBry 1912; reeve’s fig. 22a were erroneously claimed as L. f. fasciatus by clench (1932); we intend to clarify this issue in a future paper (GonzÁlez Guillén et al., in preparation). reeve’s Pl. 9 fig. 28, which is identified as A. alabaster ranG, seems like a L. f. achatinus morph arenarius sanchez roiG 1948 or L. f. achiatuinus var. romanoense jaume 1952, in error presented as a West African species. reeve’s fig. 29 is L. fasciatus var. crenatus conform his statement; his fig. 30, A. reeveana PFeiFFer, should be regarded a shell of L. f. solidulus PilsBry 1912 from Stock Island with the two peripheral bands fused, or perhaps a vari-ety of L. f. pictus without dots. Plate 10 fig. 34 is L. f. pictus reeve, fig. 35a is L. f. fasciatus as we understand it today, fig. 35c L. f. achatinus (because of the reddish columella and bigger size perhaps the var. trinidadense jaume 1952); figs. 36 a–c are L. virgineus.

A special mentioning may be made for PFeiFFer (1857), who produced a lithographical plate with two

figures of his Achatinus poeyanus, a ventral and lateral view; this taxon is currently considered as L. fasciatus by richardson (1993).

Finally, at the turn of the century, PilsBry published his treatment of this group in his monumental Manual of Conchology (PilsBry 1899: 160–175), in which he rec-ognized four full species, viz. Liguus blainianus (Poey 1853), L. poeyanus (PFeiFFer 1857), L. virgineus (lin-naeus. 1758), and L. fasciatus (müller 1774). He devel-oped a synoptic scheme to comprise the many varieties of L. fasiciatus (more details in GonzÁlez Guillén et al., in preparation). Part of his figures were copied from the original sources, while others partly they were drawn after specimens in the Philadelphia collection.

In the beginning of the 20th century the photograph-ing of shells gradually replaced other illustrating tech-niques. The first photographs of Liguus species (Fig. 8) were published by PilsBry in his treatment of the variation within Florida colonies (PilsBry 1912); however, illustrat-ing these color-full shells initially required a special tech-nique. “The original artwork was a paste-up of individual figures, which seem to have been monochrome photo-graphs that had been printed fairly pale and then hand colored (by PilsBry; he put “PilsBry pinxit” [= painted] at the bottom of each plate). This paste-up was then print-ed in color using the half-tone process, which had been around for a few years by then” (P. callomon, pers. com-mun.). A few years afterwards the first full-color photo-graphs of Liguus appeared in simPson (1920: frontispiece) (Fig. 9). That this milestone in Liguus iconography was published in a book aimed at a general public, may have contributed to their popularity with under shell collectors.

Acknowledgements

In our search for Liguus pictures and literature we had the valuable help of jonathan aBlett and kamila harPer-reekie (Natural History Museum, London, UK), Paul cal-lomon (Academy of Natural Sciences of Drexel University, Philadelphia, USA), and staff from the Naturalis Library. dolF van BruGGen (The Hague) provided help with translation of Latin texts and kindly commented on a previous version of the manuscript, which helped considerably to improve it. We thankfully mention the comments of an anonymous reviewer, who made some valuable suggestions.



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Manuscript submitted: 11.11.2013Revised manuscript accepted: 13.03.2014

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Documents

Color-full and eye-catching: an iconography of Liguus land ... · Polymita. are colorful and plentiful — at least in those days —, and were desired objects for conchologists