THE STORY OF ALCYONIUM: FROM HALCYON BIRDS TO ZOOPHYTES E Robson

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E Robson. THE STORY OF ALCYONIUM: FROM HALCYON BIRDS TO ZOOPHYTES. Vie et Milieu / Life & Environment, Observatoire Océanologique - Laboratoire Arago, 2002, pp.217-222. ￿hal-03198950￿

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HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. VIE MILIEU, 2002, 52 (4) : 217-222 From Marine Ecology to Developmental Biology In Honour of Pierre Tardent (1927-1997)

THE STORY OF ALCYONIUM: FROM HALCYON BIRDS TO ZOOPHYTES

E.A. ROBSON School of Animal & Microbial Sciences, The University of Reading, P.O. Box 228, Reading RG6 6AJ, UK

ALCYONIUM ABSTRACT. - The name of the soft coral Alcyonium is derived from the ancient ALCYONARIA Greek myth of Ceyx and Alcyone. They were transformed into halcyon birds whose HALCYONS floating nests, empty after the young hatched, were mistakenly identified with Me- ZOOPHYTE MYTHOLOGY diterranean flotsam including the remains of sessile colonies or seaweeds detached HISTORY by wave action and so variously referred to as alcyoniums. During the eighteenth century ground-breaking research on Hydra by Trembley (1744) was followed by précise descriptions of hydroids and of Alcyonium digitatum (L.) (de Jussieu 1742 and especially Ellis 1755) and the récognition that ail "zoophytes" were animais. Subclass Alcyonaria and related taxonomic terms were derived later from the name Alcyonium.

ALCYONIUM RÉSUMÉ. - Le nom du polypier charnu Alcyonium provient de l'ancien mythe grec ALCYONARIA de Ceyx et Alcyone. Ils furent transformés en alcyons, dont les nids flottants, vides ALCYONS après Péclosion des jeunes, furent identifiés par erreur parmi les débris flottants de ZOOPHYTE MYTHOLOGIE Méditerranée où se trouvaient les déchets de colonies sessiles ou d'algues déta- e HISTOIRE chées par les vagues, et donc de soi-disant alcyoniums divers. Pendant le XVIII siècle les recherches révolutionnaires sur Y Hydra de Trembley (1744) furent sui- vies par les descriptions exactes d'Hydroïdes et à'Alcyonium digitatum (L.) (de Jus- sieu 1742 et surtout Ellis 1755), et la perception de tous "zoophytes" comme animaux véritables. La sous-classe Alcyonaria (= Octocorallia) et les noms de pro- ches sous-groupements taxonomiques furent dérivés ensuite du nom Alcyonium.

The influence of Greek mythology upon biologi- Glancing back 50 years we see the post-war cal nomenclature is considérable (Thompson 1895, 1950's as trailblazing years for modem physiology, 1947) though nowadays often forgotten. As the an- for medicine and for the future molecular biology cient origins of names recède into the mists of time and genetics. Powerful computers were being de- the names become detached from them. Freed from veloped and chess players felt challenged. The the- their original context many names in use since the ory of plate tectonics was still dormant and its im- eighteenth century or earlier have survived far- plications were not generally recognized. On the reaching changes in the perspectives of science. other hand the centenary of "The Origin of Spe- The history of the name Alcyonium, briefly pre- cies" was about to be celebrated. sented here, is an example which illustrâtes how Stepping a hundred years thence to the 1850's nomenclature may evolve from myth and poetry as one finds horsepower coexisting with steam as rail- well as science. I hope Pierre Tardent would have ways and factories developed. Darwin's séminal liked this story, for he appreciated the resources of influence was as dawn still below the horizon. But history and would never underestimate what earlier new precepts of comparative anatomy and physiol- authors had discovered (Tardent 1987, 1993). ogy were already being adopted, the cell theory had The story runs from antiquity to the eighteenth gained ground and génération and development and nineteenth centuries and is set in scientific per- were beginning to be understood. Scientific knowl- spectives earlier than our own. Before telling it, an edge was increasingly enriched by travel and ex- attempt is made to recapture thèse perspectives and ploration. Scientists communicated through letters to bring less familiar modes of thought into the pic- and meetings, and published thanks to a growing ture. This account begins with thumbnail sketches number of scientific societies and journals. Their which recède century by century from modem early éducation at this time might include Latin if times to the Renaissance (Singer 1931, 1959, not Greek. Singer & Underwood 1962, Bodenheimer 1958, The century before this differed even more from Petit & Théodoridès 1962, and also Roger 1997, our own. In the 1750's Linnaeus' work shone like a Jahn 1998). beacon. It may be recalled that at this time Aris- 218 ROBSON EA totle's "Ladder of Nature" still spanned the living version of the myth is the best introduction to other world (Singer 1931, 1959) and fossils remained sources (e.g. Graves 1992, Forey 2002). puzzling. Many fine naturalists turned their atten- Alcyone, daughter of Aeolus god of the winds tion to animais and plants, sometimes minerais; and of Anarete was the wife of Ceyx, son of Luci- and they created or enriched muséums and wrote fer. They were king and queen of Trachis. Ceyx books. By 1750 microscopy had become an impor- and Alcyone found perfect happiness together but tant tool. Leeuwenhoek's sophisticated use of sim- tragedy befell them. Ceyx decided to consult an or- ple lenses for high magnification had been unique, acle and despite Alcyone's pleadings, he set sail, but for other researchers even low magnifications promising to return. His ship was wrecked in a were far better than none. The observant naturalists storm and Ceyx was drowned. Alcyone, anxious who transcribed what they saw were often pioneers with waiting, prayed for his safe return until Juno, (Trembley 1744). To an enthusiastic contemporary weary of pétitions, sent her messenger to the god of the century was remarkable for the discoveries of sleep, who dispatched Morpheus to appear in the electricity and of Hydra spp. (Le Cat 1754). At the form of Ceyx in Alcyone's dream. Alcyone sees frontiers of science knowledge was communicated her drowned husband only too clearly and awakes really fast through correspondence and meetings. It in anguish, crying out and too distraught to take was an exhilarating period even if those interested comfort. She throws herself into the sea to join him in new ideas were sometimes held back by the in- and as she does so she is transformed. Jupiter takes fluences of legend and history. Within medicine we pity on the bereft lovers and Alcyone and Ceyx are would recognize the basis of anatomy and of some turned into halcyon birds. aspects of physiology and pharmacy, although in 1750 much still lay ahead: most of the basis of Halcyons are kingfishers, but to relate the myth modem chemistry, the contributions to physiology as transcribed to détails of the birds known to Aris- of von Haller and J Mûller, and advances in the un- totle or other ancient observers is not straightfor- derstanding of disease due to Jenner, Pasteur and ward. One might think of terns or gulls or else gan- Lister. Latin served as a lingua franca for spécial nets but the halcyon had a tapered beak and was texts, and was an accessible médium. brightly coloured, with blue head and back and a reddish breast (Thompson 1895, 1910, Belon 1555: To step across another hundred years towards 218). None of the présent species of kingfisher cor- 1650 is to expérience in a différent era the full vig- responds to halcyons in nesting habits or distribu- our of scientific discovery and discussion. Charters tion. were granted to the Royal Society in 1662 and to the French Académie des Sciences in 1666. The gi- There is more to the myth. Halcyon birds make a ants on whose shoulders we stand include Newton, nest which floats unharmed upon a tranquil sea for Boyle, Hooke, Malpighi, Haies and many others seven days before and seven days after the winter whose achievements still influence today's young solstice. The winds drop as Aeolus restrains them. scientist. By 1650 entrainment in Latin and Greek The female lays her eggs and broods, the young scholarship had become less germane to science birds are hatched and they fly away as the calm and thus from the beginning, the Royal Society of halcyon days end. Then Aeolus releases the unruly London's Transactions, for example, were pub- winds. The substance of the nest is uncertain, in- lished in English. cluding sea foam and perhaps fishbones, but care- fully fashioned with the beak to be seamless and The records of yet earlier authors in the six- unbreakable (e.g. Aldrovandi 1603: 497). teenth century are of significant interest here as will be seen. Works on natural history, whether er- Sailors' legends follow: occasional sightings of udite compilations (e.g. Gesner 1555, Aldrovandi "halcyon birds" or calm midwinter spells in the 1603) or new (Belon 1555) provided detailed re- Gulf of Corinth confirm the myth. Halcyon birds in views of ancient literature, both Greek and Latin. flight are propitious as they bring good weather. A Scholars of ail persuasions used Latin themselves female flying alone is heard to call repeatedly, as if and were steeped in Homer and Ovid and so famil- in distress. Maritime legends about halcyons in- iar with classical mythology. cluding the médicinal properties of sea foam per- sist well into the sixteenth century (e.g. Belon The subject of this narrative are the soft corals 1555: 219). représentée! by Alcyonium digitatum (L.), a British Ovid's transcriptions of earlier sources have coastal species commonly known as dead man's stood the test of centuries and have influenced al- fïngers. It is in the family Alcyonaceae, order most every Western writer (Hughes 1997: vii). The Alcyonacea, subclass Alcyonaria (Hickson 1930) myth of Ceyx and Alcyone retains ail its vitality. It otherwise Octocorallia (Manuel 1981, Fabricius & has inspired poetry, opéra and works of art (Reid Alderslade 2001). 1993). The phrase "halcyon days" is still a literary The source of the name Alcyonium is the Greek or political metaphor indicating a fortunate period myth of Ceyx and Alcyone, which gave rise also to of untroubled calm (Simpson & Weiner 1989). Hal- the metaphor "halcyon days". Ovid's well known cyon birds have featured in heraldry and also as THE STORY OF ALCYONIUM 219

emblems (Giovio 1574, Scorza 1981). Fig. 1 is a Distinctions between animais and plants were device from Giovio (1562) which shows halcyon sharpened by Abraham Trembley's séminal experi- birds on their floating nest during the days of calm ments on freshwater Hydra spp. (Trembley 1744). weather. The rubric and accompanying verse (Mr Thèse were greeted with amazement if not disbe- CG Wagstaff kindly furnished this translation) are lief. That animais which captured prey and walked innovative: "The halcyon birds know the elected about could bud, and even regenerate like plants, time for the sea not to harm their nests or their one type being green besides, caused a great stir, eggs. Unhappy the man who knows not how to wait and an upheaval of previously held ideas (e.g. to give effect to his design". Giovio's device for Réaumur 1742, Dawson 1987). Trembley's beauti- the Fieschis refers to their strategy for taking up ful work is still as good as new and marks the rise arms in revenge, which proved successful. It was of expérimental biology (Baker 1952, Lenhoff & an âge when both myth and observation were sig- Lenhoff 1986, Tardent 1987). Trembley sent hydra nifïcant. Paulo Giovio (1483-1552) and the natural- to de Réaumur, who in 1741 confirmed his conclu- ist Pierre Belon (1517-1564) were near contempo- sions. It was he who suggested the term "polype" raries. for hydra by analogy with the Polypi (i.e. cephalo- pods) of ancient authors, whose eight arms capture The term Alcyonium sprang from the coexis- prey (Réaumur 1742: li-lxxx; Baker 1952). tence of myth, legend and reality. Halcyon birds' nests of sea foam interwoven with fishbones might The influence of Trembley's research was im- later reappear among the floating flotsam and jet- médiate and widespread. In 1755 Ellis refers to sam of Mediterranean coasts. Poorly specified ob- "the Freshwater Polype whose extraordinary prop- jects were called alcyonium or alcyoneum from an- erties have been so fully enquired by the ingenious cient times (e.g. Pliny: see Thompson 1895). Light Mr Trembley". Twenty years later it is common weight or of spongy texture, perhaps compound in knowledge. "Every one knows, that the common origin, they might include soft corals, sponges, as- polype sends out its young from its side, like cidians, bryozoans and encrusted algae detached by buds..." (Ellis 1776). wave action (e.g. Ellis 1755, Ellis & Solander Linnaeus (1758) placed A. digitatum in his 1786, Johnston 1838). Some of the sponges re- Zoophyta nonetheless. Thèse were composite ani- ferred to as alcyoniums had ancient médical uses, mais with plant-like stems in which he included sea an aspect of sea foam which influenced later hal- pens, gorgonians, hydroids and Hydra. He placed cyon legends (e.g. Thompson 1895, 1947, Belon corals under Lithophyta. Ray (1724), however, had 1555: 219). By the end of the sixteenth century the included Alcyonium digitatum and several misapprehension that the several kinds of lithophytes and zoophytes of Linnaeus (e.g. corals) alcyonium were halcyons' nests had faded away. under aquatic plants (Plantae Submarinae). Ellis The soft coral Alcyonium digitatum appears in (1755) used Ray's Latin description of A. the lOth édition of Linnaeus' "Systema Naturae", digitatum. His English rendering for Fig. 2 is the species described as Alcyonium ramoso- "Alcyonium of a soft fleshy Nature, with its Sur- digitatum molle by John Ellis (1755) and shown in face full of Stars". Fig 2. It was well known from flatfish grounds Ellis used low magnification (a microscope by (Ray 1724). English fishermen called it dead man's John Cuff with single lenses) to great advantage hand, or dead man's toes, or cow-paps (Johnston and he was besides an acute and thoughtful ob- 1838). The Mediterranean species Alcyonium server. He saw living polyps of hydroids and later palmatum (Pallas 1766) was called manus marina those of Alcyonium digitatum (Ellis 1754, 1755, (the sea hand) (Gesner 1558:619, Aldrovandi 1763). In the sponge Halichondria panicea, whose 1606:593). Ancient Greek sponge divers would texture resembled that of A. digitatum, there were probably have seen it. no polyps. Instead, oscula opened and closed and Early in the eighteenth century the boundaries water passed through them (Ellis 1763, 1765). He between animais and plants were by no means clear was in no doubt that ail of thèse spécimens were (Robson 1975). In Aristotle's "Ladder of Nature" animal in nature. animais were higher than plants, and minerais were Comparable but less detailed observations were inanimate. The idea of a zoophyte, in the sensé of made by Bernard de Jussieu in 1741. Familiar with an animal with vegetable attributes, began with Hydra spp. and the earlier view of Peysonnel Aristotle. Later naturalists observed that there were (1752) that corals were animais, he visited the aquatic plants with stony parts (lithophytes). If the north coast of France and upon studying Tubularia natural world showed every aspect of indivisa and Alcyonium digitatum he had reached intergradation lithophytes and zoophytes offered the same conclusions (de Jussieu 1742, Allman catégories which avoided taxonomic boundary 1871). problems and were consistent with contemporary outlook. The word "zoophyte" was widely used un- Although Linnaeus (1758) incorporated his dis- til the late nineteenth century, by then in a descrip- coveries Ellis did not think that zoophytes had af- tive sensé (Simpson & Weiner 1989). finities with plants. His careful observations re- 220 ROBSON EA

Fig. 1. - Device by Paolo Giovio (1562) for Sinibaldo and Ottobuono Fieschi of Genoa, showing halcyon birds on their nest under a clear sky. During seven days before and after the winter solstice (the "halcyon days") it floats unharmed upon a calm sea. Giovio (1574) had in mind Pliny's account of the myth. The birds were to be shown in bright colours (azuré, red, white, green and yellow). The rubric in French "We know the right time" and the accompanying Italian verse (see text) refer to the Fieschi's strategy for revenging themselves upon their adversaries, which proved success- ful. Photo: Warburg Institute.

solved uncertainties about the status of those with which we so much admire in a natural historian." branching skeletons such as gorgonians (Ellis (Ellis 1776). 1776). As he was still called "the father of English Zoophytology" in 1883 (Simpson & Weiner 1989) During the nineteenth century the range of soft his concluding views on myth and natural history corals and related Anthozoa (sea fans, sea pens, red may be quoted: coral and others) became better known: knowledge of local marine species increased as did the collec- "... although they grow in a branched form, they tion of spécimens from much further afield. Sev- are no more allied to vegetables than they are to the eral higher taxonomic terms derived from the name ramified configurations of sal ammoniac; to the él- Alcyonium were introduced, such as the subclass égant branched figures in the Mocha and other Alcyonaria (Fabricius & Alderslade 2001: 8). gems, called dendrites; to the arbor Dianae, or the Thèse aspects may be traced in a splendid history arborescent figures of the Cornish native copper: of researches on Anthozoa by Carlgren (1903). For consequently, that animal life does not dépend on earliest use of the term Alcyonaria see Dana bodies growing according to a certain external (1846), Milne-Edwards (1857) and Hickson form. Hence it appears, that this metamorphosis of (1930). For inspired illustrations of soft corals see a plant to an animal is a flowery expression, and in Saville Kent (1893) and Fabricius & Alderslade my opinion, better suited to the poetical fancy of an (2001). Hickson's fïrst impressions are still vivid OVID, than to that précise method of describing (Hickson 1889): "This was my first introduction to THE STORY OF ALCYONIUM 221

Fig. 2. - Alcyonium digitatum (L.), the "Alcyonium of a soft fleshy Nature with its Surface full of Stars", from John Ellis (1755). The spécimen attachée! to an oyster shell shown at a was collected by fishermen near Whitstable, Kent. At A the surface papillae magnified by a lens are seen with 8-pointed stars from which polyps émerge. As shown in A2, their 8 tentacles are fringed. Photo: University Photographie Unit, Reading. a coral reef... I was not prepared to find such bril- Baker JR 1952. Abraham Trembley of Geneva scientist liancy and variety of colour..." "... it was impossi- and philosopher 1710-1784. Edward Arnold, London, ble to put one's foot down on anything save living 259 p. zoophytes..." and while out on low spring tides Belon P du M 1555. Histoire de la Nature des Oyseaux, "...watching... the slow and graceful waving avec leurs descriptions, et naifs portraicts retirez du movements of the Alcyonarian polypes...". naturel: escrite en sept livres, etc. Gilles Corrozet, Pa- ris, 382 p. ACKNOWLEDGEMENTS. - I would like to thank Library Bodenheimer FS 1958. The History of Biology. William Staff at the British Library, The Institute of Classical Dawson & Sons, London, 465 p. Studies, the Linnean Society, the Natural History Mu- séum, the Royal Collège of Surgeons, the Royal Society, Carlgren O 1903. Coelenterata (Hohlthiere). Anthozoa. the University of Reading, the Wellcome Library, and HG Bronns Klassen und Ordnungen der Thierreichs. the Librarians and Photocollection Staff at the Warburg Bd II, Abt 2, Buch 3, Lief 1-6. CF Winter, Leipzig, Institute for their help, and Mrs M Barratt for typing the 176 p. manuscript. Dana JD 1846. Structure and Classification of Zoophy- tes. Lea & Blanchard, Philadelphia, 132 p. Dawson VP 1987. Nature's Enigma: the problem of the polyp in the letters of Bonnet, Trembley and Réau- REFERENCES mur. Mem Amer Phil Soc Philadelphia 1 74: 266 p. Ellis J 1754. A Letter from Mr John Ellis, F.R.S. to Mr Allman GJ 1871. A monograph of the Gymnoblastic or Peter Collinson, F.R.S. concerning the animal Life of Tubularian Hydroids. The Ray Society, London, those Corallines that look like minute Trees, and 450 p. grow upon Oysters and Fucus's ail round the Sea- Aldrovandi U 1603. Ornithologiae Tomus Tertius aepos- coast of this Kingdom. Phil Trans 48: 627-633. tremus. Lib XIX - XX. Bologna, 560 p. Ellis J 1755. An Essay towards a Natural History of the Aldrovandi U 1606. De Reliquis Animalibus exsangui- Corallines and other Marine Productions of the like bus libri quattuor... de Mollibus, Crustaceis, Testa- Kind, Commonly found on the Coasts of Great Bri- ceis et Zoophytis. Bellagamba, Bologna, 593 p. tain and Ireland, etc. London, 103 p. 222 ROBSON EA

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