Typification of Cannabis sativa L.
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ABSTRACT
The name Cannabis saliva L., published by Carl Linnaeus in 1753 in his Species plantarum, the internationally accepted basis for modern botanical nomenclature, is the first legitimate scientific name for the hemp grown in Europe where it had been extensively cultivated for many centuries previously. The cannabis material which Linnaeus had for study is fortunately represented by two good specimens in the Clifford Herbarium of the British Museum of Natural History. Although Linnaeus gave "India" as the country of origin of the species, he based his description on the hemp grown in northern Europe in 1737, which he knew in a living state and that had been described at length by Rabelais, in 1545, under the fictitious name "Pantagruelion."
The possibility that the genus Cannabis (Cannabaceae) comprises more than one species, as believed by Zhukovsky (1962) and other Russian botanists and as noted by Tutin et al. in Flora Europaea 1:67 (1964), or consists of one variable species divisible on fruit characters into several subspecies with differing chemical properties, has made it essential to examine the typification of the name Cannabis saliva so as to remove in advance any nomenclatural uncertainty which may otherwise come about if, for taxonomic reasons, the Linnaean epithet saliva has to be restricted to one member of the group.
The name Cannabis saliva L., having been published by Carl Linnaeus in his Species plantarum 2:1027 (1753), the internationally accepted starting point for modern botanical nomenclature, is the first legitimate scientific name for the hemp which was grown in Europe in the 18th Century. Here it had been extensively cultivated for many centuries, as is evident from both historical and palynological evidence (summarized by H. Godwin in 1967), being grown primarily for its tough fibres providing cordage and clothing but also for its oily seeds; fortunately, during the period of its maximum use in Europe, the narcotic properties of its resin were unknown there.
The flowers of hemp are either male (staminate) or female (pistillate). Normally an individual plant bears either male or female flowers but not both. Male and female individuals differ in appearance and longevity, the males having conspicuous loose few-leaved inflorescences and dying earlier than the females, which have compact more leafy inflorescences with much less conspicuous flowers.
Growers of hemp have probably always been familiar with the differences between male and female plants and have long distinguished them as such in a metaphorical manner completely opposed to their biological nature. According to Lefranc (1905), Antoine Rabelais, the father of François Rabelais (c. 1494-1553), grew much hemp on his property at Cinais, southwest of Chinon (Indre et Loire), and young Rabelais probably helped in its cultivation. Rabelais certainly knew everything known then about the character and cultivation of hemp; three chapters of his Le tiers livre des Faictz et Diaz héroiques du noble Pantagruel (1546) are devoted to l'herbe nommée Pantagruelion, which is simply hemp. Rabelais here duly mentioned its sexuality:
Et, comme en plusieurs plantes sont deux sexes, masle et femelle, ce que voyons es lauriers, palmes, chesnes, heouses, asphodele, mandragore, fougere, agaric, aristolochie, cypres, terebynthe, pouliot, peone, et aultres, aussi en ceste herbe y a masle qui ne porte fleur aulcune, mais abonde en semence, et femelle, qui foisonne en petits fleurs blanchastres, inutiles, et ne porte semence qui vaille, et comme est des aultres semblables, ha la feuille plus large, mains dure que le masle, et ne croist en pareille haulteur (Livre 3, chap. 49).
Sir Thomas Urquhart in his 1693 translation came closer to the original than he usually did, being swept along by his exuberant love of words which Rabelais would have appreciated, when he rendered this into English as follows:
And as in diverse plants and trees there are two sexes, male and female, which is perceptible in laurels, palms, cypresses, oaks, holmes [i.e. holmoaks], the daffodil [i.e. asphodel], mandrake, fern, the agaric [i.e. mushroom], birthwort, turpentine, pennyroyal, peony, rose of the mount and many other such like, even so in this herb there is a male which beareth no flower at all, yet it is very copious of and abundant in seed. There is likewise in it a female, which hath great store and plenty of whitish flowers, serviceable to little or no purposes, nor doth it carry in it seed of any worth at all, at least comparable to that of the male. It hath also a larger leaf and much softer than that of the male, nor doth it altogether grow to so great a height.
The seed-bearing hemp called "male" here is, of course, the female plant and the sterile hemp here called "female" is really the male.
Most pre-Linnaean botanical authors, except Ray and Morison, applied the terms mas (male) and foemina (female) in the same metaphorical way as Rabelais, without any concept of true sexuality in plants comparable to that of animals. Thus, of two kinds, usually distinct species, the more robust or more vigorous or more useful one, especially if having larger leaves or harder wood, was designated "male" and the inferior one "female." Hence, the names Cannabis saliva and C. mas, as used by D'Aléchamps, Dodoens and C. Bauhin, refer to female individuals of hemp; and the names C. erratica, C. foemina and C. sterilis refer to male individuals. The name Cannabis saliva, which Linnaeus used as a specific name covering both sexes, applied originally only to female individuals. This kind of usage died slowly. As late as 1884, Saint-Lager noted, in his erudite "Remarques historiques sur les mots 'plantes males' et 'plantes femelles'," that farmers in the Rhône basin were still calling pistillate plants of hemp "chanvre mile" and staminate plants "chanvre femelle," because the pistillate plants remained green and robust after the weaker staminate plants had withered, their function as pollinators fulfilled.
In the same manner, C. Bauhin designated the useful female hop-bearing plant of Humulus lupulus as Humulus mas and the unproductive male as H. foemina.
The difference between male and female plants of hemp necessitates two periods of harvesting. Thus, Philip Miller in his Gardeners dictionary, 8th ed. (1768), recorded that in the east of England,
the first season for pulling the Hemp, is usually about the middle of August, when they begin to pull what they call the Fimble Hemp, which is the male plants.... These male plants begin to decay soon after they have shed their farina. The second pulling is soon after Michaelmas, when the seeds are ripe: this is usually called Karle Hemp, it is the female plants, which were left at the time the male were pulled.
The fruit is a small nut, i.e. it has a single seed tightly covered by the hardened wall of the ovary, and is enclosed within a sheathing hairy bract with abundant resin glands which presumably developed in the wild as a protection for the fruit against insects, like the glandular trichomes of other plants (cf. D. A. Levin 1973). The distinctions, which have been made between the taxa known as C. saliva, C. indica and C. ruderalis, relate to characteristics of the fruit; male plants seemingly provide no diagnostic features; hence for typification a pistillate specimen would be preferable to a staminate one on taxonomic as well as historic grounds.
Linnaeus's protologue in the Species plantarum 2: 1027 (1753) is as follows:
1. CANNABIS sativa
Cannabis foliis digitatis. Hon. cliff 457.
Hort. ups. 297. Mat. med. 457.
Dalib. paris. 300. Roy. lugdb. 221. Cannabis sativa. Bauh. pin. 320. Cannabis mas. Dalech. hist. 497. Cannabis erratica. Bauh. pin. 320. d Cannabis femina. Dalech. hist. 497. d Habitat in India.
Several matters in this protologue call for comment. In genera with several species, Linnaeus provided concise diagnostic phrase-names enabling the species thereby to be distinguished, e.g. Hippophae fouis lanceolatis and Hippophaé follis ovatis for H. Rhamnoides and H. cana-densis. Such phrase-names were comparative; they contrasted specific features. In a genus with only one species, such as Cannabis, no such diagnostic phrase was required and would indeed have been illogical, since obviously the one and only species could not be contrasted with itself.
Typification of the generic name of such a monotypic genus is essentially the same as typification of the specific name; the nomenclatural type of the one must be the nomenclatural type of the other. Hence, the generic name Cannabis L. and the specific name Cannabis saliva L. must be permanently associated with the same element. The Species plantarum citations of literature begin with Linnaeus' own Hortus Cliffortianus (1738), where fuller synonymy will be found; the other citations likewise refer to plants cultivated in Europe. He used the terms mas and fern/na and the signs du and for male and female plants in a purely biological sense and sorted his synonyms accordingly. Knowing hemp only as a cultivated plant in Europe, he evidently assumed that it must have been introduced from elsewhere, presumably Asia; he had earlier identified with this the male plant figured under the name "Kalengi-cansjava" in Rheede, Hortus Malabaricus 10: t. 60 (1690) and, with some doubt, the female plant figured there in t. 61(1690); on this evidence, it would seem, he stated "Habitat in India." These Rheede illustrations were later cited by Lamarck under his Cannabis indica when he separated that as a species distinct from C. sativa.
Linaeus' account of Cannabis saliva in the Species plantarum (1753) is to be associated with the description of the genus Cannabis in his Genera plantarum, 5th ed., 453, no. 988 (1754), as stated in the International Code of Botanical Nomenclature art. 13, note 3 (1972). This description is as follows:
988. CANNABIS* Tourne!. 308
* Mas
CAL. Perianthuim quinquepartitum: foliolis oblongis, acuminatoobtusis,
concavis.
COR. nulla.
STAM. Filamenta quinque, capillaria, brevissima, Antherae oblongae, tetragonae.
* Femina
CAL. Perianthium monophyllum. oblongum, acuminatum, latere altero longitudinaliter dehiscens, persistens.
COR. nulla.
PIST. Germen minimum. Styli duo, subulati, longi. Stigmata acuta.
PER. minimum. Calyx arcte clausus.
SEM. Nux globoso-depressa, bivalvis.
The asterisk in the heading CANNABIS* here, as in the first edition, indicates that Linnaeus had based his account on living material, i.e. on plants cultivated in Sweden or Holland. This 1754 description comes, however, unchanged from the first edition of the Genera plantarum 304, no. 749 (1737) published at Leyden, when Linnaeus had charge of Cliffords' richly stocked garden at Hartekamp. That work, dealing with the genera, and his Hortus Chffortianus (1738), dealing with the species, have the same close association as the 1754 Genera plantarum has with the 1753 Species plantarum. Thus, his principal reference under Cannabis in the 1753 Species plantarum is to the Hortus Chffortianus 457, which, in turn, refers to the 1737 Genera plantarum no. 749.
In the Hortus Cliffortianus, Linnaeus provided a short diagnosis, Cannabis fouis digitatis, to distinguish the true hemp from a then imperfectly known plant diagnosed there as Cannabis fouis pinnatis, but named Datisca cannabina in the first edition of the Species plantarum. In short, Linnaeus' concept of Cannabis saliva in 1753 is identical with that of his Cannabis folds digitatis of 1738. Just as John Ray had earlier distinguished functionally male individuals as Cannabis sativa "mas s. sterilis" and female individuals as Cannabis sativa "foemina s. fendis," so Linnaeus likewise distinguished male and female individuals, allocating preLinnaean synonyms to each. The material of Cannabis which Linnaeus had for study when preparing the Genera plantarum (1737) and Hortus Cliffortianus (1738) is fortunately represented in the Clifford Herbarium, Hortus siccus Cliffortianus, in the Department of Botany, British Museum (Natural History), London, by two good specimens, one (A) male (Plate 1.), the other (B) female (Plate 2.). Either is available for designation as lectotype. Since, however, the major characters for taxonomic division in cannabis come from fruiting material, the Hortus siccus Cliffortianus fruiting specimen (p. 457 Cannabis no. 1, B) of Cannabis sativa L. is here designated as the lectotype. This specimen represents C. sativa as currently commonly accepted. The fruit is about 5 mm. long, 3.5 mm. broad.
If Linnaeus had provided in 1753 a new diagnosis for Cannabis sativa or had modified in 1754 the generic description of Cannabis published in 1737 on the basis of later material — as he did for some other species and genera — then it would be judicious to select a lectotype from this material influencing his final concept of these. In fact, however, he did neither. Hence, as indicated above, the lectotype has to be taken from the earlier material on which his unchanged concepts were based. From this standpoint, the two specimens under Cannabis in his herbarium at the Linnaean Society of London have only a subsidiary relevance, because they in no way affected his publications. They are, however, of interest on account of their Linnaean association. Linnaean Herbarium specimen 1177.2, illustrated in Joyce & Curry, The botany and chemistry of Cannabis 22 (1970), is a pistillate plant, with fewer than the usual numbers of leaflets, which are narrowly lanceolate, long acuminate and Sharply serrate. It has no epithet but is numbered "1" in Linnaeus' hand.
Linnaeus began to draft his Species plantarum long before he devised his method of consistent binominal nomenclature for species; even in 1748, he had not devised binomials for the whole vegetable kingdom; hence the most convenient method of arranging and designating his herbarium specimens was to number the species in each folder according to the numbered species entries in his manuscript Species plantarum. When, a few years after 1753, he began to prepare a second edition of the Species plantarum, with changed numbering of specific entries, he ceased to number his specimens but added instead the specific epithet introduced in that work. Thus, a numeral corresponding to an entry in the first edition of the Species plantarum is a valuable indication that Linnaeus possessed this specimen in 1753 or acquired it soon afterwards.
Hence, the numbered pistillate specimen with leaflets characteristic of European hemp, Linn. Herb. 1177.2, can be assumed to have been in his hands at this time if not much earlier. The other specimen, Linn. Herb. 1177.1, illustrated in Joyce & Curry, Botany and chemistry of Cannabis 21 (1970), is of very different aspect. It is a staminate plant with much shorter and broader almost obtuse more coarsely serrate leaflets. It has no number but is labelled "sativa" in Linnaeus' hand. Thus, this specimen, in no way typical of Cannabis saliva as commonly accepted, can safely be assumed to have come into Linnaeus' possession later than 1753.
The two Hortus Cliffortianus specimens belong to the old cultivated hemp stock of northern Europe. This is represented by another contemporary herbarium specimen in the British Museum (Natural History) which was grown in the Chelsea Physic Garden and presented in 1740 to the Royal Society of London under the number 908; for a discussion of the history and nomenclatural importance of these Chelsea specimens, see Steam (1972). There are also specimens scattered through the herbaria assembled by Sir Hans Sloane (1660-1753) and now in the British Museum (Natural History): vol. 39, fol. 2 (c. 1660). vol. 83, fol. 161 (L. Plukenet, 1642-1706), vol. 85, fol. 62 (G. Bonnivert, II. 1673-1703), vol. 91, fol. 47 (Plukenet), vol. 117, fol. 2 (A. Buddle, 1660-1715), vol. 167, fol. 393 (G. London, d. 1713), vol. 321, fol. 236 (H. Boerhaave, 1668-1738); see J. E. Dandy (1958).
SUMMARY
Although Linnaeus, when publishing the name Cannabis sativa in 1753, gave "India" as the country of origin of the species, he based his original description on the hemp grown in northern Europe in 1737, which he knew in a living state; this hemp belonged to the long-cultivated European stock which Rabelais had described at length in 1545 under the fictitious name "Pantagruelion." Linnaeus, like his predecessor Ray, correctly distinguished the staminate individuals as "male" and the pistillate and fruiting individuals as "female." Most prepLinnaean authors, on the general masculine assumption that males were superior or more robust or more useful than females, metaphorically designated the relatively useless male individuals as "female" and the fruit-bearing female ones as "male." A female (pistillate) cultivated specimen in the Clifford Herbarium at the British Museum (Natural History), London, is taken as lectotype of the name Cannabis sativa L.
REFERENCES
DANDY, J. E., editor
1958 The Sloane herbarium. London: Trustees of the British Museum.
GODWIN, H.
1967 The ancient cultivation of hemp. Antiquity 41:42-50,137p138.
1967 Pollen-analytic evidence for the cultivation of Cannabis in England. Rev. Palaeobot. & Palynol. 4:71-80.
JOYCE, C. R. B., S. H. CURRY, editors.
1970 The botany and chemistry of Cannabis. London: J. & A. Churchill.
LEFRANC, A.
1905 Pantagruelion et Chenevreaux. Rev. Études Rabelaissennes 3:402-403.
LEVIN, D. A.
1973 The role of trichomes in plant defense. Quart. J. Biol. 48:3-15.
SAINTpLAGER, J. B.
1884 Remarques historiques sur les mots 'plantes males' et 'plantes femelles.' Ann. Soc. Bot. Lyon 11 (1883):1-48.
SCHULTES, R. E.
1970 "Random thoughts and queries on the botany of Cannabis," in The botany and chemistry of Cannabis. Edited by C. R. B. Joyce and S. H. Curry, 11-38. London: J. &. A. Churchill.
STEARN, W. T.
1970 "The Cannabis plant: botanical characteristics," in The botany and chemistry of Cannabis. Edited by C. R. B. Joyce and S. H. Curry, 1-10. London: J. &. A. Churchill.
1972 Philip Miller and the plants from the Chelsea Physic Garden presented to the Royal Society of London, 1723-1796. Bot. Soc. Edinburgh Trans. 41:293-307.
ZHUKOVSKY, P. M.
1962 Cultivated plants and their wild relatives. Translated by P. S. Hudson. Farnham Royal (Commonwealth Agricultural Bureau).
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