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"That illustrious pair of true friends, Scipio and Laelius, joined one to another by the bond of affection and also by partnership in all virtues, just as they followed the path of active life with equal tread, so did they find relief together in mental relaxations. For it is agreed that they used to pick up shells and pebbles [conchulas et umbilicos] wandering on the beaches of Caieta and Laurentum."

Valerius Maximus, Memorable Doings and Sayings (VIII.8) from the anecdote related by Cicero, De Oratore (II.22)

Beginning in the eighteenth-century, there was a craze for seashells, especially exotic rarities and newly discovered specimens, that rivaled the earlier mania for tulips. This conchylomania (from the Latin concha, "shellfish") derived from the colonial trade and exploration of the Dutch East India Company, which dominated the market in China and Indonesia. Porcelain and spices were imported, as well as shells that were eagerly sought by wealthy collectors.

In 1796, for example, a shell from the collection of the Dutch naturalist Pierre Lyonet sold at auction for 299 guilders; another, a Conus cedonulli that was two inches long sold for 273 guilders. Although it is not known how much his Conus gloriamaris realized, Lyonet had paid 120 guilders for it, three times the selling price of Johannes Vermeer's Woman in Blue Reading a Letter, which had sold at auction for 43 guilders.

The earliest specimen of C. gloriamaris belonged to a Dutch collector, as did most examples, and first was mentioned in a 1757 sales catalog, where it is listed as Gloria maris. It is this rather modest (92 mm) and damaged shell (a growth line runs almost its entire length) that was the holotype (the definitive type specimen) for the scientific description of the species by Chemnitz in 1777.

By 1865, ten were thought to exist, all from Dutch cabinets, except for two live specimens that had been found by an English collector in 1837 on a reef in the Philippines, which later was thought to have been destroyed. Two dozen were known by 1957, one of which sold that year for $2000, and perhaps twice that number when Dance's book was published in 1966. Then, only two years later, the habitat of C. gloriamaris in the Solomon Islands was discovered and it finally became affordable (above).

 Woman in Blue Reading a Letter, c.1632-1663 (Rijksmuseum, Amsterdam)

Lyonet's collection (described in the sales catalog as du célèbre cabinet de Coquilles de feu Pierre Lyonet) sold at auction on April 21, 1796, six-and-a-half years after his death. Dance relates the prices paid for two of the shells (C. cedonulli and the Nautile Vitré) and then compares what they realized to the selling price of two paintings by Jan Steen, which sold for 50 guilders; two by Franz Hals, which sold for approximately 1 and 10 guilders, respectively; and the Woman in Blue Reading a Letter by Vermeer, which sold for 43 guilders. Given the juxtaposition of the two paragraphs (and the fact that the primary source for Dance was in French), the assumption has been that the shells and paintings sold at the same time.

In fact, as the provenance of Woman in Blue indicates, the painting had been auctioned by Lyonet's estate five years earlier, on April 11, 1791. It was then, too, that the paintings by Steen (of which Lyonet owned four) and Hals were sold. The following is the description of Vermeer's masterpiece from the sales catalog: "In an inner room one sees a young lady dressed in a blue satin jacket standing at a table on which lie a small box, pearls, and papers; she seems to be reading a letter with much attention; on the white wall hangs a large map, filled in with other accessories; this piece is beautifully and naturally treated on canvas, by the Delft van der Meer, the pleasing light and dark lend it a fine appearance as is commonly characteristic of the works of this famous Master."

This conflation of the two auctions invariably occurs in the literature, most recently in an article in Smithsonian. A Christie's catalog (September 23, 2004) has the Vermeer painting selling in 1791 for 41 guilders and includes the Nautile Vitré.

Prices no doubt would have been higher for Lyonet's shells were it not for the political turmoil in Europe at the end of the eighteenth century. Louis XVI had been guillotined in 1793 and the Dutch republic abolished by the French in 1795. The next year, a month before the auction, Napoleon invaded Italy.

One-hundred years earlier, when the last voyage of discovery by the Dutch East India Company was taking place, there had been a remarkable sale of 21 Vermeer paintings inherited by the son-in-law of his most important patron. The catalog description allows many of them to be identified. At that auction in 1696, View of Delft sold for 200 fl; The Milkmaid, 175 fl; Woman Holding a Balance, 155 fl; The Music Lesson, 80 fl; The Concert, 73 fl; and Woman Writing a Letter with Her Maid, 63 fl. The Girl with a Turban (The Girl with a Pearl Earring), perhaps Vermeer's most iconic painting, sold at auction in 1881 for only two guilders (plus a 30% premium), the picture too damaged and dirty to have revealed the signature of the artist.

The Concert was stolen in 1990 (having been acquired almost a century earlier), and a C. gloriamaris that had been presented to the American Museum of Natural History almost thirty years before was stolen in 1951. Only two other specimens then were known to be in the United States, and an article at the time excitedly had announced the gift and plans to display it. Neither the painting nor the shell have been recovered.

Because Isabella Stewart Gardner's will stipulated that the exhibit remain on view exactly as she had arranged it, with no picture to be removed or replaced, the empty frame still is presented as a silent reproach to the crime.

  The shell that sold for 299 guilders, seven times the price realized by Vermeer's painting, was Carinaria cristata ("Glassy Nautilus") or, as it then was called, the Nautile Vitré. Lyonet owned the earliest recorded specimen, which seems to have been only one of two examples. A small shell, looking like a glassine pouch, it sold for 575 fl when put at auction again three decades later, more than twice the sum paid for a C. gloriamaris from the same collection.

The Portland Vase (British Museum)

One of the most notorious collectors of the time was the second Duchess of Portland, whose purchase of shells and other objects was so excessive that, ten years prior to the Lyonet auction, everything of hers had to be sold, including this eponymous first-century BC Roman blue cameo glass vase, which she had purchased in 1784 from Sir William Hamilton. At auction two years later, it realized £1,029.

There are only thirty-six paintings by Vermeer, who died when he was forty-three. A Young Woman Seated at the Virginal, the first one to come to market since 1921, sold at Sotheby's in 2004 for almost £16.25 million ($30 million), more than five times its pre-sale estimate. This small canvas was the last of his paintings still in private hands but after a decade of analysis, the attribution to Vermeer finally allowed it to go to auction.

Given the notorious forgeries of Han van Meegeren (who managed to sell a fake Vermeer to Hermann Göring and confessed only to avoid an accusation of collaboration with the Nazis), it is perhaps understandable that the painting would have been suspect. In 1927, Meegeren's The Lacemaker had been declared "a genuine, perfect, and very characteristic work of Jan Vermeer of Delft" when presented for authentication to Wilhelm von Bode, the first curator of what is now the Bode Museum (Berlin), where the forgery first was exhibited. It was not so much the imitation of Vermeer's style that convinced the experts but Meegern's ability to capture what they themselves, in that time and place, imagined that style to be. The picture, in other words, was thought to be modern, rather than typical of the era in which it was created. And this may explain why The Lacemaker now looks so unprepossessing, in spite of its pointillés, the small bright dabs of paint looking like sequined points of light that are characteristic of Vermeer.

Later that year, the painting was sold to Andrew Mellon, who also purchased The Smiling Girl, another forgery by Meegeren (although it is possible that both were by his mentor Theo van Wijngaarden). It, too, had been pronounced by Bode as a "characteristic work" of Vermeer, and the figure in the portrait did look very similar to The Lacemaker—but smiling. At Mellon's death in 1937, both pictures were donated to the National Gallery of Art (Washington, DC), which he founded and where the pictures were exhibited. They now are in storage and not on view (to some consternation at the time, surprisingly). One cannot feel too much sympathy for the defrauded Mellon, who had the Internal Revenue Service prepare his tax returns and whose personal wealth more than doubled during his tenure as Secretary of the Treasurer under President Calvin Coolidge.

So expensive were exotic shells from the South Seas that they were regarded as investments (which actually was the name of one such specimen) and celebrated in seventeenth-century Holland pronkstilleven (from the Middle Dutch pronken, "to show off"; from which the Middle English pranken derived and from it, "prank" and "prance"). These ostentatious or sumptuous still lifes of luxurious and precious objects included both tulips and shells, the attraction of one being its ephemerality, the other its immutability. There not only was the satisfaction of collecting these works of nature and the aesthetic appeal of their variety and beauty, but the status such rare objects bestowed upon the owner. Wealthy collectors identified with their collections, sought to preserve them and be known for them—and be remembered as a result.

The most famous of the Dutch shell painters was Balthasar van der Ast, whose painting above is in the Gemäldegalerie Alte Meister (Old Masters Picture Gallery), Dresden. Two of the shells, which are so common now, are the Tiger Cowrie (Cypraea tigris) and Marble Cone (Conus marmoreus), which Rembrandt etched in 1650, more than ten years later. Curiously, it was drawn as he observed it (and not reversed) so that, when the paper was pulled, the image did not reproduce correctly. The aperture appears on the left (as it would in a sinistral shell) instead of the right.

Just before his death in 1789, Lyonet made quite specific arrangements for his shell collection, which he requested remain complete and in Holland. It therefore was to be offered first to various societies; if after a month, there were no offers, it then was to be advertised in the better known French journals. Only if the collection remained unsold after four months was it to be cataloged and put up for auctionwhich it was.

When another collector had died almost a century and a half earlier, the executors of the estate found his collection of tulips neatly stored in labeled drawers and boxes, so many in number that the notary recorded thirty-eight pages in the inventory. Every shell was counted, a total of almost twenty-four hundred. As Lyonet later would do, precise conditions were specified for their disposal. Each of three executors was to have a separate key to the chest that contained the shells so that no-one could sell or tamper with them unless all three men were present. It also was stipulated that the collection remain intact for four years. Only then could the shells be sold individually.

A recent study published by the National Academy of Sciences offers a fascinating computer simulation of how mollusks construct and pigment their shells. A network of nerve fibers in the mantle, the tongue-like protrusion that wraps around the leading edge of the shell, is connected with secretory cells that stimulate the deposit of calcium carbonate, proteins, and other organic material, including pigmented compounds that are the nitrogenous waste products of metabolism. Different rates of calcium carbonate secretion determine the shape of the spiraling shell and the rhythmic secretion of pigment, the pattern. What is intriguing is that this pattern, laid down at the edge of the growing shell, itself regulates the incremental layering of shell material, the mantle sensing the calcium carbonate layer deposited the previous day to create a new layer the next.

For example, when a pigment-secreting cell inhibits the secretion of pigment by surrounding cells and the same pattern is repeated, a stripe is displayed. Pigments secreted one day that are inhibited for a few days from further secretions result in an on-off pattern of bands parallel to the growing edge. And when pigment inhibits secretion at one site but excites it at another, the pigment is secreted laterally on successive days, which produces zigzags, chevrons, and other patterns.

As beautiful as these shell patterns are, they seemingly are irrelevant to the mollusk, which typically hides during the day under rocks or in the sand and hunts at night. Too, the shell may be obscured by a rough translucent periostracum (and the algae attached to it) and the pattern not visible even to a potential mate. Rather than display or camouflage, the deposit of pigment helps the mollusk build a shell of the right shape by getting its mantle in register, so to speak. The pattern of the growing shell, itself, is only an epiphenomenon, a secondary record of neurosecretory activity. And yet, it is no less beautiful for that. Unwhorled, the shell can be imagined to look like the traces of an electroencephalogram. But instead of a graphic record of human neural activity, it is the neurological record of the mollusk as recorded in the pattern of its pigmented calcium carbonate.

Equally varied are the venoms of cone shells. Each species has evolved a unique concoction of toxic proteins, which are delivered by a sharp harpoon-like tooth in the proboscis that darts out and spears the victim, paralyzing it. Because the mollusk cannot chase its prey or risk injury when it is caught, these conotoxins have evolved to immobilize almost immediately. It is this ability that has made the study of toxic cones so important in pharmacological research—and, like the patterns of their shells, such fascinating creatures.

"A true practical philosopher, dear old Mussard lived without a care in the world in a very agreeable house he had had built for himself and in a very pretty garden he had planted with his own hands. While excavating under the terraces in this garden he had found some fossilized shells, and had found them in such quantities that his exalted imagination began to see shells everywhere in nature, and so that he came in the end to believe that the universe was made up of shells and the remains of shells and that the whole earth consisted of nothing but crushed shells. Wholly absorbed in this subject and in his singular discoveries, he became so excited by his ideas that they would in the end have combined themselves in his fevered brain into some system, which is to say some folly, if, very fortunately for his reason, but most unfortunately for his friends, to whom he was very dear and who found his house the most agreeable of sanctuaries, death had not snatched him from them through the strangest and most cruel of illnesses."

Rousseau, Confessions (II.8)

The French term conchyliologie was introduced by d'Argenville in the first edition of his book on shells (1742) and defined in the second edition (1757). Later in the paragraph, Rousseau uses the word conchyliomanie to describe his cousin's mad preoccupation with shells. The English equivalent "conchylomania" (or concholomania) has not yet found its way into the revised Oxford English Dictionary, however, where the entry for "conchology" still derives from the first edition (1891). There, the Elements of Conchology: or, An Introduction to the Knowledge of Shells (1776) by the British naturalist Emanuel Mendes da Costa is cited as the first use of "conchology" in English. The word seems to have appeared several years earlier, however, in Conchology, or A Natural History of Shells (1771) purported to have been co-authored by da Costa.

This was at a time when mollusks were studied almost exclusively in reference to their shells. "Malacology" (malacologie in French, first used in 1814) is the branch of zoology that deals with the study of mollusks and, although the distinction between the two words can be muddled, conchology tends to refer in popular usage to a study of the shell, malacology to the snail that produces it.

The Oxford Latin Dictionary translates concha as "mollusk" or "shellfish"; so Plautus in his play Rudens ("The Rope," 297) speaks of fishermen returning with conchas. Later, it is the shell from which Venus herself was said to have been born (704). More particularly, the word also denotes the murex and oyster, as well as the shell of the mollusk, mother-of-pearl, and pearls. Although sometimes translated as cockle or mussel (as in the Smithsonian article), Plautus, in describing the fishermen's catch, uses musculus to denote mussels (298).

Mollusks comprise the phylum Mollusca and are classified according to the characteristics of their shells. The word derives from the French mollusque (1783), "soft-bodied" (Latin mollis, "soft" as in "mollify") and denotes several classes of snails that are protected by their exoskeletons. Gastropoda such as Conus, for example, have a single (univalve) spiraling shell, bivalves such as oysters and mussels have two valves or shells, hinged together. "Mollusk" tends to be the American spelling of the word, "mollusc," the British.

There does not seem to a collective noun for shells (what has been called a noun of multitude, such as an "exaltation of larks"). But, if there were, a "Caligula of shells" might be appropriate, given the anecdote related by Suetonius, who records the emperor's planned expedition against Britain in AD 40, which did not advance beyond the English Channel.

"Finally, as if he intended to bring the war to an end, he drew up a line of battle on the shore of the Ocean, arranging his ballistas and other artillery; and when no one knew or could imagine what he was going to do, he suddenly bade them gather shells and fill their helmets and the folds of their gowns, calling them 'spoils from the Ocean, due to the Capitol and Palatine'" (Life, XLVI).

References: Shell Collecting: An Illustrated History (1966) by S. Peter Dance (the standard reference for the history of shell collecting); Tulipmania: Money, Honor, and Knowledge in the Dutch Golden Age (2007) by Anne Goldgar; A Prosperous Past: The Sumptuous Still Life in the Netherlands, 1600-1700 (1988) by Sam Segal [exhibition catalog]; "Mad about Shells" (August 2009) by Richard Conniff, Smithsonian Magazine, 40(5), 44-51; "The Case History of a Rare Shell: Conus Gloriamaris Chemnktz 1777" (September 1970) by E. R. Cross and Rutn Fair, Hawaiian Shell News, 18(9), 1, 3-8; "High Hopes to Fallen Dreams" (May 1971) by Alan Solem, Field Museum of Natural History: Bulletin 42(5); "Shell Sales. II" (1942) by J. R. le B. Tomlin, Journal of Molluscan Studies, 25(1), 25-33; "Type Specimens and Identity of the Described Species of Conus II. The Species Described by Solander, Chemnitz, Born, and Lightfoot Between 1766 and 1786" (1964) by Alan J. Kohn, Zoological Journal of the Linnean Society, 45(304), 151-167; "The Neural Origins of Shell Structure and Pattern in Aquatic Mollusks" (2009) by Alistair Boettigera, Bard Ermentroutb, and George Ostera, Proceedings of the National Academy of Sciences, 106(16), 6837-6842; "The 'Glory of the Sea'" (1923) by Roy Waldo Miner, Natural History, 23(4), 325-328; "Diversity of Conus Neuropeptides" (1990) by B. M. Olivera et al., Science, 249(4966), 257-263; Vermeer of Delft: Complete Edition of the Paintings (1978) by Albert Blankert; The Man Who Made Vermeers (2008) by Jonathan Lopez; Pierre Lyonet, 1706-1789 (1962) by W. H. Van Seters (in French); Jean-Jacques Rousseau: Confessions (2008) translated by Angela Scholar (Oxford World's Classics); Valerius Maximus: Memorable Doings and Sayings (2000) translated by D. R. Shackleton Bailey (Loeb Classical Library).

The article in Smithsonian is one of very few to appear in the popular literature. The classic is "Shells Take You Over World Horizons" (July 1949) by Rutherford Platt, The National Geographic, 96(1), 33-84 (with 41 color photographs), written at a time when the golden cowrie (Cypraea aurantium) was worn only by tribal chiefs and could not be exported from Fiji. Another important article is "The Magic Lure of Sea Shells" (March 1969) by Paul A. Zahl and Victor R. Boswell, The National Geographic Magazine, 135(3), 386-429.

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