BULLETIN OF MISCELLANEOUS INFORMATION No. 2; 1939
Royal Botanic Gardens, Kew

AMARYLLIS AND HIPPEASTRUM
J. R. Sealy
I. INTRODUCTION

In the first edition of his "Species Plantarum" (1, 292-293: 1753) Linnaeus placed nine species in the genus Amaryllis L., and these nine species belong to as many different genera according to our modern concepts of the genera of Amaryllidaceae. As a result of this original wide concept of the limits of Amaryllis, still further diverse elements were included within it during the years that followed the publication of Linnaeus's work, and by the beginning of the nineteenth century it had come to include about fifty species representing at least a dozen different genera. Some of the original nine species of Amaryllis were referable to certain genera (Brunsvigia Heister, Sprekelia Heister, Imhofia Heister, Atamasco Adanson, and Sternbergia Waldst. & Kit.) that had been proposed during the period between 1753 and 1805, but these genera either remained unknown to students of the family or were not recognized by them, and the genus Amaryllis was generally accepted as being very polymorphic.

The task of sorting the diverse elements that constituted this Amaryllis, sensu latiore, into homogeneous groups and defining them as distinct genera was first attempted in 1819 by Herbert (Bot. Mag. sub t. 2113). Two years later he dealt with the problem in more detail, correcting his previous work in certain respects, and extending it to include all the genera of Amaryllidaceae, and this work, published under the title "An Appendix" (actually to vol. 6 of the Botanical Register), forms the basis of all the succeeding taxonomy of the family. In the "Appendix" Herbert restricted the name Amaryllis to the Cape plant now so well-known to botanists and gardeners alike as Amaryllis Belladonna, and distributed the other diverse elements of Amaryllis among other genera, some of which he established for their reception

*This name was based on Amaryllis equestris Linn. fil. ex Aiton, Hort. Kew. 1, 417 (1789), which, however, is antedated by Amaryllis puniceus Lamarck, Encycl. Method, 1, 122 (1783). The correct name for the species under Hippeastrum is accordingly H. puniceum (Lam.) Urban, Symb. Antill. 4, 151 (1903), but for the purposes of this paper the better-known name H. equestre is retained. [Note: Lamarck identified his Amaryllis punicea as A. Belladonna of Linnaeus. Furthermore, we may note Amaryllis Belladonna described in the 1768 edition of Miller's Dictionary is the American plant.]

For some years students of the Amaryllidaceae have known that the Cape plant to which Herbert applied the name Amaryllis Belladonna was not, apparently, the plant that Linnaeus meant when he first gave the name to one of the species of Amaryllis in the first edition of his Species Plantarum (1753). Under the name Amaryllis Belladonna Linnaeus cited certain synonyms, which evidently all refer to the American plant now known as Hippeastrum equestre (Linn. fil.) Herb.,* and not to the Cape Belladonna at all; in fact Linnaeus gave the habitat of his species as "in Caribaeis, Barbados, Surinama." The matter was mentioned by the late Dr. Stapf in a footnote on page 3 of his article on XCrinodonna Corsii in the Botanical Magazine, t. 9162 (1929), but nothing was published about the problem until last year, when an article by Dr. Uphof appeared in "Herbertia" (1938, pp. 100-109, tt. 107-109). Uphof comes to the conclusion that Amaryllis Belladonna L. is the correct name for Hippeastrum equestre (Linn. fil.) Herbert, that the genus Hippeastrum Herbert must be renamed Amaryllis L., and that the Cape Belladonna must receive the name Callicore rosea Link. These conclusions seem reasonable at first sight, but on closer examination it appeared that the complexities of the case had not been fully appreciated by Uphof, and that it would be necessary to reconsider it in more detail and from a different angle before one could judge whether or not his conclusions were correct.

The case was accordingly fully re-investigated and the results are given below. The problems concerned are much more involved than would appear from Uphof's account, and form another example of the difficulties so often experienced when attempts are made to identify and typify Linnaean genera and species, and illustrate the necessity of caution in dealing with such problems.

Whilst taking entire responsibility for the views expressed in this paper, I gratefully acknowledge the help given to me, by way of advice and discussion, by Messrs. J. Ardagh and J. E. Dandy of the British Museum (Natural History) and by Mr. Spencer Savage of the Linnean Society. Dr. T. A. Sprague has been kind enough to go over the MS. in detail with me, and I am greatly indebted to him for the many suggestions he has made and the advice he has given.

II. THE IDENTITY AND TYPIFICATION OF
AMARYLLIS BELLADONNA L.

The name dates from Linnaeus, Species Plantarum, 1, 292 (1753), where the species is defined as "Amaryllis spatha multiflora, corollis campanulatis aequalibus, genitalibus declinatis" and a reference is given to Linnaeus, Hortus Cliffortianus, 135 (1737) where that phrase name is used for the first time. In addition Linnaeus cites certain synonyms, with references to earlier authors, and gives the habitat of the species as "in Caribaeis, Barbados, Surinama." The references, synonyms, and habitat apply, as stated above, to the American plant we know as Hippeastrum equestre, and therefore at first sight it seems clear that Amaryllis Belladonna L. was actually Hippeastrum equestre (Linn. f.) Herb. Three points, however, suggest caution in accepting this conclusion:

* For example, Miller, Gard. Dict. ed. 1 (1731). Actually the Cape Belladonna was in cultivation in Italy as early as 1633, fide Ferrari, De Florum Cultura, 116-117, c.c (1633). In the Italian version of that work (Flora overo Cultura di Fiori, 118: 1638) the common name is given as "Donna bella.") The American plant was in cultivation in Italy prior to 1625, according to Aldinus, Hortus Farnesianus (1625). Ferrari also described it.
(1) three authors of Linnaeus's own time, Philip Miller, John Miller, and the younger Linnaeus, actually applied the name Amaryllis Belladonna to the Cape plant known under that name to-day;1 (2) we know from contemporary literature*2 that the Cape Belladonna was a well-known garden-plant in Linnaeus's day, [As was the American Belladonna, according to Sloane and Hill.] and it seems incredible that Linnaeus would not have known of it, and if he knew the plant, it is only reasonable to suppose that he would have included it in his "Species Plantarum" [Linnaeus neglected the far more common Rosa damascena Mill.]; (3) as already mentioned, Linnaeus diagnosed his A. Belladonna with a phrase taken from his "Hortus Cliffortianus," a work which deals with the identification and classification of plants grown in the garden of Georg Clifford at Hartecamp, Holland, and of specimens in Clifford's herbarium, so that we can say that Linnaeus must have seen either a dried specimen or living plants (or both) of his Amaryllis Belladonna, and that this species was not based merely on the synonyms he quotes. This fact was apparently unnoticed by Uphof, but at once raises the question, was that specimen or plant really the American Hippeastrum equestre, or could it have been the Cape Belladonna?

Further investigation has brought to light the evidence detailed below:—

(1) In the "Viridarium Cliffortianum," Linnaeus gave a list of all the living plants in the Clifford garden during the years 1735, 1736 and 1737, and among them is the "Amaryllis spatha multiflora, corollis campanulatis aequalibus, genitalibus declinatis," the plant which he subsequently named Amaryllis Belladonna. Linnaeus therefore certainly knew a living plant of this species.

(2) In the Clifford herbarium, now at the British Museum (Natural History), London, there is a specimen immediately recognizable as the Cape Belladonna. The specimen bears no name or identification, and therefore there is no ground for stating either that it is the basis of Amaryllis Belladonna, or that it is the plant which Linnaeus knew in the Clifford garden, for the specimen may have been added to the herbarium after Linnaeus had left Holland. On the other hand, the absence of data on the sheet suggests that this specimen was grown in Clifford's garden, and there is a possibility that the species was growing there, or that the specimen was in Clifford's herbarium, when Linnaeus was writing the "Hortus Cliffortianus," and it is therefore possible that the specimen represents one of the species described in that work. It is obviously referable to the genus Amaryllis and can be referred only to the second species, the one which in 1753 Linnaeus named Amaryllis Belladonna.

*A. sarniensis (Nerine), A. zeylanica (Crinum), A. longifolia (usually placed in Ammocharis), A. orientalis (Brunsvigia), and A. guttata (Boophone).

(3) The Cape Belladonna and the American Hippeastrum equestre are markedly dissimilar plants, and if Linnaeus had known the latter in cultivation and had based his diagnosis of Amaryllis Belladonna upon it, one would expect the fact to be evident from his diagnosis. The latter reads: "Amaryllis spatha multiflora, corollis campanulatis aequalibus, genitalibus declinatis" and there are two points in it which are not in keeping with Hippeastrum equestre. The first is "spatha multiflora," which is scarcely descriptive of a plant 2-4 flowers, even if used as a contrast to species "spatha uniflora". Had Linnaeus been describing Hippeastrum equestre, it is surely more likely that he would have written "spatha pauciflora," and it should be noticed that all the other species of Amaryllis* described as "multiflora" in the "Species Plantarum," ed. 1, really are many-flowered.3 On the other hand, in the second edition of the "Species Plantarum," Linnaeus accepted Miller's description of A. Reginae as "multiflora," though this species is generally 2-4-flowered. It might therefore be said that "multiflora" should be taken as meaning "more than one-flowered" in contrast to the "uniflora" of other species. It should be remembered, however, that in the case of A. Reginae Linnaeus was merely recording Miller's work, whereas for A. Belladonna the diagnosis was original and written by Linnaeus himself.4

The second point concerns the "corollis campanulatis." No one who knew living plants (and it must not be forgotten that Linnaeus knew his Amaryllis spatha multiflora...etc. in life) or even good figures of Hippeastrum equestre, could describe the flowers as campanulate.5 The flower has a most characteristic shape, as shown in figure 1 (actually taken from one of the works cited by Linnaeus), and I suggest that it is exceedingly unlikely that Linnaeus would have described it as "corollis campanulatis." On the other hand, both the "spatha multiflora" and the "corollis campanulatis" are understandable if Linnaeus was dealing with the Cape Belladonna, for in that plant the stem bears 10 or 12 flowers, which, as shown in figure 2, would be accepted as campanulate as that term was used by Linnaeus.

(4) In the "Hortus Cliffortianus," under Amaryllis, Linnaeus wrote:—"Lilio narcissus vocabulum est consarcinatum, quod rejicio. Flores hujus generis eximii sunt, nescio num 2-da parem habeat, hinc Bellae donnae dictae plures; Bella donna virgilii, amaryllis dicta, nomine transiit in proverbium de omni grato, et de secunda specie apud Hortulanos quosdam, quae cum et radice amara sit, pro Amarella Amaryllis dicatur."

This may be translated:—"Lilio-narcissus is a patched-together designation which I reject. The flowers of this genus are very beautiful—I do not know that the second species has an equal—hence many are called Belladonna. The Belladonna of Virgil was called Amaryllis and this name has passed into proverb for any beautiful thing, and has been used among the gardeners for the second species, which although it has a bitter root, is called Amaryllis in place of Amarella."

The "second species" to which Linnaeus referred in this note is the one he named A. Belladonna in 1753, and two points stand out at once. First that the species had such beautiful flowers that Linnaeus doubted if it had an equal, and second that the species was at least fairly well-known in cultivation since it had already received the name Amaryllis in gardens. Both remarks apply to the Cape Belladonna—no one would dispute that its flowers are exceedingly beautiful, and it was certainly well-known in gardens in Linnaeus's time (see next paragraph)—but one could scarcely claim that Hippeastrum equestre had no equal in beauty of flowers, and it was certainly not well-known in cultivation in Linnaeus's day, in fact it was very rare. [Nehrling, who came upon a bed of H. equestre in Florida wrote of "a shimmering mass of beautiful, fiery red flowers. ... I was speechless from astonishment and inspiration."]

(5) The Cape Belladonna was in cultivation in Italy in 1633. It was included by Morison in his "Plantarum Historiae" (1680), and by Miller in his "Gardener's Dictionary" from the first edition (1731) onwards; it was apparently introduced into this country at an early date and became well-known. Hippeastrum equestre may also have been in cultivation in England in Miller's time, for his Lilio-Narcissus no. 3, the West Indian Red Lily (Gard. Dict. Ed. 1-4: 1731-43), stated to be very rare in England, may have been this species. It should be noted that the West Indian Red Lily of Ed. 6 and Ed. 7 (1752-59) was another species, H. Reginae and that this had been introduced in 1728.6 The Cape Belladonna was growing at Leyden in 1681-1686 and is included in Hermann's "Catalogous" (1687), where, incidentally, it is called Lilium africanum...etc. It was at Leyden when Boerhave prepared his "Index Alter" (1720), and in that work there is also a reference to the American Hippeastrum equestre.

*Royen was assisted by Linnaeus in his re-arrangement of the Leyden garden—see Daydon Jackson, "Linnaeus," 163 (1923)
Only one species is mentioned, however, by Royen in his account of the Leyden garden (1740) and that species was recorded under the Linnaean phrase-name* for Amaryllis Belladonna, with synonyms belonging to the American plant. It is scarcely likely that two such different plants as the Cape Belladonna and the American Hippeastrum would be identified with one another in a garden, and hence it is most likely that one or the other had disappeared from the Leyden garden by 1740. Of the two, the American plant, which has to be grown under glass and requires careful treatment, is more likely to have disappeared than the hardy Cape plant.7 It is therefore much more likely that the plant recorded by Royen was the Cape Belladonna. All the available evidence, in fact, points to the Cape Belladonna being well known in gardens in Linnaeus's day, whereas the American Hippeastrum equestre was rare. [Note: Miller (1731) wrote that the Cape Belladonnas were "very rare", which Sealy takes to mean "well known". By contrast, the American Belladonna was described as "very rare", which Sealy implies meant almost non-existent. The fact that a plant has a common or vulgar name does not imply that it is common.]

(6) Philip Miller, in his "Figures of Plants" (1755) figured the Cape Belladonna under the name Amaryllis spatha multiflora...etc. of the "Hortus Cliffortianus," and in the second edition of the "Species Plantarum," Linnaeus cited Miller's figure as an illustration of his Amaryllis Belladonna.8 Miller stated that the plant was well known in [Italian] gardens as the Belladonna lily, and that it had been identified by Sir Hans Sloane with a plant which he had obtained in Barbados and which he had called Lilio-narcissus polyanthos, flore incarnato, fundo ex luteo albescente (Sloane, Cat. 115, Hist. I. 244).9 Miller goes on to say, however, that from all he can learn there are two species of "lily" in the West Indies, one white-flowered and evidently a Pancratium, the other red-flowered and certainly not the Belladonna lily. In the next plate of his "Figures," Miller gives an illustration of the plant known in English gardens as the "Mexican Lily," which, he says, had been named Lilium Reginae by Dr. James Douglass, "because it was in full Beauty on the First of March, which was the late Queen's Birth-day." This plant is the one known to-day as Hippeastrum Reginae. With it Miller identified the Lilium Americanum puniceo flore, Belladonna dictum described and figured by Hermann in his "Paradisus Batavus" 194 (1698), but he adds "Doctor Herman says it came from the Caribbee Islands; but all the roots which I have received from those Islands, by the Title Red Lily, are of a different sort from this." In this statement Miller was perfectly correct, for the Red Lily of the West Indies is H. equestre. The mistake that Miller made was in identifying Hermann's plant with H. Reginae, for Hermann's plant is unmistakably H. equestre. Now Linnaeus included Miller's "Mexican Lily" as a new species of Amaryllis in the second edition of his "Species Plantarum" and named it A. Reginae, quoting, as a synonym, the Hermann reference already mentioned. This means that under A. Reginae his synonym referred to another plant, Hippeastrum equestre, whilst the references in the same work under A. Belladonna likewise belonged to H. equestre. It may be noted that in one of the works in the synonymy of A. Belladonna, namely, Merian's "Metamorphosis," the plant described and figured is identified with the Hermann plant [and Sloane's], and also that in the "Hortus Cliffortianus" Linnaeus had actually included the Hermann reference in the synonymy of the plant he afterwards names A. Belladonna. The conclusion seems to be that Linnaeus did not critically examine the literature, and that he had no clear idea of the identity of the plants described and figured in the works cited. In fact, it is most likely that the synonyms cited by Linnaeus in the "Hortus Cliffortianus" represent "book-work" hastily compiled owing to the great pressure under which that work was written.10

In the second edition of the "Species Plantarum" Linnaeus emended his diagnosis of Amaryllis Belladonna by adding "ungue reflexis" to the original "corollis campanulatis aequalibus." This was evidently put in the better to distinguish it from A. Reginae, for under the latter he says it differs from A. Belladonna by the petals being undulate at the margin and not recurved into a claw ("nec ad ungues recurvatis"). Now Linnaeus did not get this additional character from Miller, for the latter does not mention it, nor is it shown in his plate, and we must therefore conclude that Linnaeus derived it from a plant or specimen which he accepted as his own Amaryllis Belladonna. [This is a deliberate mistranslation. "Ungues" or "claws" are the bases of petals, a usage that can be traced back at least as far as Pliny the Elder. Cape Belladonna petals are "unguiculate" or "clawed" at the tips, which is not what Linnaeus was describing: "...ad ungues recurvatis" means bent at the base.] This recurving [incurving!] of the apex of the outer perianth-segments in the shape of a claw is characteristic of the Cape Belladonna, but does not occur, so far as I am aware, in Hippeastrum equestre. In the latter the apices may be shortly apiculate or minutely cucullate but can scarcely be described as "recurved in a claw." The claw-like apices are very clearly shown in John Miller's excellent figure of the Cape Belladonna in his "Illustration of the Sexual System of Linnaeus," t. 18, part of which is reproduced in figure 2. This work was known to Linnaeus, who formed a high opinion of it, but the plate of Amaryllis Belladonna did not appear until 1770 (date on plate), that is, long after the second edition of the "Species Plantarum." [In the 1779 edition, J. Miller mentioned 1777 as the year of the first edition. He only stated that the common name was BELLADONNA, in agreement with the Gardeners Dictionary discussion of Amaryllis regina.] It would seem that some time between 1753 and 1762 Linnaeus became acquainted with Miller's "Figures of Plants," and that before accepting Miller's "Mexican Lily" as a new species of Amaryllis, he examined a plant or specimen of his own A. Belladonna to see how the new species differed from it, and that he noted the claw-like apex of the outer segments. This would mean that Linnaeus was perfectly aware of the identity of his A. Belladonna, and that that species was the Cape plant. It may be of interest to add that in his own copy of the "Species Plantarum," ed. 2, Linnaeus has written in a short description for A. Belladonna, which reads as follows: "Corolla regularis campanulata statura Hemerocallidis, incarnata, fundo viridi albicante. Petala 3 exteriora apice ungue intus retro spectante. Petala 3 superiora basi ciliata. [Beard pads!] Stamina declinata rubra. Antheris albidis. Pistillum rubrum apice albo. Caulis teres, parum compressus. Bulbis viridis." This evidently applies to the Cape Belladonna, and together with the work published in ed. 2 of the "Species Plantarum," shows conclusively11 that Linnaeus did accept the Cape plant as his A. Belladonna some time between 1753 and 1762; but it does not necessarily mean that he based the species on the Cape plant when he first described it in 1737. The fact that Linnaeus did accept the Cape plant as A. Belladonna, however, in conjunction with the fact that the 1737 description and reference to the outstanding beauty of the flowers both apply to the Cape plant, seems to justify the conclusion that the species was based on the Cape plant. It may be mentioned that Trew (Hort. Amoen. Fl. Imag. Ed. Seligmann, 1, p. N2. verso, t. 12) also described the Cape plant under the phrase that Linnaeus used in 1737, and figured it as "Belladonna." The title-page of volume I is dated 1750, and if the work had been written prior to that date, it would mean that even before 1753 the Cape plant had been recognized as the species described us in 1737. However, on the page of Trew cited is a reference to another work the date of which is given as 1766, so that Trew's work must have appeared during or after that year.12

*Reprinted in Linné, Amoen. Acad. Ed. Schreberus, VIII. pp. 249-267 (1785)

(7) In 1775 a thesis, prepared under Linnaeus's direction by one of his pupils, Jacobus Alm, was published under the title "Plantae Surinamensis."* This thesis deals with the identification of a collection of Surinam plants and among them is an Amaryllis which is dealt with thus:—"39. Amaryllis 98. dubia. Mer. surin. t. 22. Corolla basi laciniarum barbata." That is to say, that no. 98 was an Amaryllis whose specific identification was doubtful but which was [similar to] the plant figured by Merian. The latter, of course, is the American Hippeastrum equestre, and, since Merian's work was actually cited by Linnaeus under Amaryllis Belladonna in the "Species Plantarum," one would normally have expected that Alm would have given this name for the Amaryllis no. 98 of the collection. That he did not do so seems to imply that either he or Linnaeus had become aware that the American plant was not A. Belladonna.13

(8) The younger Linnaeus14 prepared a MS. on certain plants, including a number of Amaryllidaceae, which remained unpublished and is now in the possession of the Linnean Society of London. This MS. recently formed the subject of an article by Mr. Spencer Savage in Herbertia, 1937, 92-99, in which he showed that the MS. was written about 1781-83, and that it formed the source of the diagnoses of various plants which were published in Aiton's "Hortus Kewensis" (1789) and there attributed to "Linn. fil.," the diagnoses concerned having been sent to Aiton by Linnaeus fil. himself (see Aiton's Preface, p. vi). Among the plants dealt with are those with which we are concerned, and it is instructive to notice that the younger Linnaeus completely clears up the confusion which surrounds Amaryllis Belladonna and A. Reginae. He does this by restricting the name A. Belladonna to the Cape Belladonna (as may be easily demonstrated by the references he gives), by restricting the name A. Reginae to the plant figured by Miller, and by establishing a new species, A. equestris, to cover the American plant which Linnaeus had placed in the synonymy of A. Belladonna (by citing the works of Sloane, Seba and Merian under that species), and which Miller had misidentified with A. Reginae (when he cited Hermann, Par. Bat. 194 as a synonym of the latter). Incidentally, the habitat of A. Belladonna was not given in Aiton's "Hortus Kewensis," but it is evident from Savage's account (p. 99) that the younger Linnaeus knew that it came from the Cape, and wrote in the MS. "Habitat in Africa Australi" Cap: Bona Spei." The name Amaryllis Belladonna was also restricted to the Cape plant by L'Héritier in his Sertum Anglicum, 11-13 (1788). [Wrong! L'Héritier included plants from four distinct genera under his Belladonna. Three of his references are to American plants: Lilio-narcissus rubens indicus. Ald. farn. 83. t.82., Lilio-narcissus jacobaeus phoeniceus indicus polyanthos. Barrel. rar. 69. t.1036., A. spathâ multiflorâ, corollis campanulatis aequalibus, genitalibus declinatis. Mill. ic. 15. t.23. He also noted that A. chilensis was the same color as reginae and Belladonna, which rules out the Cape Plant.] Moreover, L'Héritier quoted much of the earlier literature concerning the plant, and gave its home as the "Cape of Good Hope," a piece of information which may have been derived from one of the works cited, namely, Petiver, Gazophylacii Nat. Art. 9, t. 85, fig. 5, et Cat. Class. Sec. Vol. p. 4, no. 497 (1764). The Sloane, Seba and Merian references, which Linnaeus senior had placed under Amaryllis Belladonna, were transferred by L'Héritier to A. Reginae. L'Héritier obviously recognized these references were to the same plant as the one figured and described by Hermann, but he apparently did not realize that they all represented a species distinct from A. Reginae.15

The evidence from the foregoing discussion may be conveniently summarized thus:—

(a) Linnaeus knew a living plant of his Amaryllis Belladonna in Clifford's garden;
(b) The Cape Belladonna was a well-known garden plant [in Italy] in Linnaeus's day; it is most unlikely that it was unknown to Linnaeus, and if he knew the plant he would certainly have accounted for it in his "Species Plantarum"; [he did in the second edition] Hippeastrum equestre was rare in cultivation; [grown in France by 1603 (Vallet), in Italy before 1625 (Aldinus), in England in 1659 (Hamner), and was always common in English-controlled Jamaica.]
(c) There is a specimen of the Cape Belladonna in the Clifford Herbarium, but no specimen of Hippeastrum equestre;
(d) Linnaeus especially mentions his Amaryllis Belladonna as an outstandingly beautiful species; it was apparently reasonably well-known in cultivation, since Linnaeus mentions the name given to it in gardens: these remarks are applicable to the Cape plant, but scarcely to Hippeastrum equestre.
(e) Linnaeus's original diagnosis of his Amaryllis Belladonna fits the Cape plant better in two respects; ("spatha multiflora" and "corollis campanulatis") than it does Hippeastrum equestre.
(f) In the second edition of his "Species Plantarum," Linnaeus cited a figure of the Cape plant as an illustration of Amaryllis Belladonna [Linnaeus referred to Plate 23—the Scarlet Belladonna.] and added to the diagnosis a character ("corollis...ungues recurvatis") present in the Cape Belladonna but not in Hippeastrum equestre. Moreover, Linnaeus must have seen this character in a specimen or living plant, for it is not mentioned in the literature, nor shown in any of the illustrations cited by him. Furthermore, Linnaeus wrote a description of his Amaryllis Belladonna in his own copy of "Species Plantarum" ed. 2, and that description is of the Cape Belladonna [He also mentioned the beard pads.] This all shows that Linnaeus accepted the Cape plant as his A. Belladonna. [It shows that Linnaeus was not referring to a preserved specimen.]
(g) Under A. Reginae Linnaeus, following Miller, erroneously cited as a synonym the Lilium Americanum...etc. of Hermann, which he himself had cited under A. Belladonna in 1737, and which is the same plant (Hippeastrum equestre) to which his synonyms under A. Belladonna belong. This indicates that Linnaeus had not examined the literature carefully. [It indicates that Linnaeus got his information from Miller's 1760 Figures where Hermann's phrase-name is mistakenly associated with the plate of the Cape Belladonna.]
(h) In 1775 the Merian reference, given in synonymy of A. Belladonna by Linnaeus in 1737 and 1753, was cited in a thesis prepared under Linnaeus's direction, as an Amaryllis whose identity was doubtful.
(i) Three botanists of Linnaeus's own times, Philip Miller, L'Héritier, and the younger Linnaeus, all agree in identifying Amaryllis Belladonna L. as the Cape plant, and excluding from it the American Hippeastrum equestre. [L'Héritier excluded very little from his Amaryllis Belladonna. He included at least four species of perhaps as many genera. Philip Miller (1768) distinguished the Cape plant as A. regina. John Miller (1779) only mentioned that the common name of this plant was "Belladonna". He had worked for J. Miller, and contributed to the 1768 edition of Gardeners Dictionary.]

Taken as a whole, this evidence indicates that Linnaeus knew the Cape Belladonna, that he almost certainly based his Amaryllis Belladonna on it, but confused it with the literature relating to Hippeastrum equestre.

If this view is rejected, we are left with the alternative that the name Amaryllis Belladonna was based only on the American plant Hippeastrum equestre, known to Linnaeus in cultivation and from the literature. The difficulties that arise when one attempts to reconcile this alternative with the evidence detailed above, will be readily discernible.

Now regarding the identification of Amaryllis Belladonna L., it seems that the four following views are possible:—

(1) We can reject entirely the suggestion that the Cape Belladonna formed a part of Amaryllis Belladonna L. in 1753, and say that only Hippeastrum equestre is concerned. This is Uphof's conclusion.
(2) We may consider that Linnaeus based Amaryllis Belladonna on the literature he cited, i.e., on Hippeastrum equestre, that he misidentified with it the Cape Belladonna known to him as a garden plant, and took the characters for the diagnosis, at least in part, from the garden plant.
(3) We can consider that Linnaeus based Amaryllis Belladonna on the Cape Belladonna known to him as a garden plant, and misidentified it with the literature belonging to Hippeastrum equestre.
(4) We may agree that Linnaeus confused the Cape Belladonna and the literature concerning Hippeastrum equestre under Amaryllis Belladonna but consider that there is not sufficient evidence to show that either species has a better claim than the other to the name Amaryllis Belladonna L.

With regard to these four possibilities I would remark as follows:—
(1) In my opinion the evidence as a whole, and particularly that under (b), (d), and (e), and to a less extent under (c), (f) and (h) makes this view so improbable that it must be rejected.
(2) and (3) These being alternatives may be considered together. It seems much more likely that Linnaeus would have had chiefly in mind the garden plant from which he took his diagnosis, and would have misidentified with it the literature connected with another species, rather than vice-versa. The diagnosis is so incompatible with the species represented by the literature, that there is no justification for giving the literature precedence as the basis for the name Amaryllis Belladonna. Consequently I think that (3) is the more reasonable view. As to the type of the name A. Belladonna, this will be the plant from which Linnaeus took his diagnosis, or specimen thereof if one exists. The specimen of the Cape Belladonna in the Clifford Herbarium may actually be the type, but unfortunately this cannot be proved. However, in the absence of evidence to the contrary this specimen may be accepted as the working type.
(4) In my opinion the evidence points to view (3) being preferable to this, and consequently I accept (3) and reject (4). However, this view might be accepted by other workers, in which case a lectotype of the name A. Belladonna would have to be chosen. As the "Regulations for determining types," mentioned in the International Rules of Botanical Nomenclature ed. 3 (1935), have not yet appeared, the choice of lectotype should be governed by the general principles upon which the Rules are based. The first essential point in nomenclature is to aim at fixity of names (Art. 4), and in the absence of a relevant rule established custom must be followed (Art. 5). The first author to restrict the name A. Belladonna to one of its two elements and definitely exclude the other, was L'Héritier, who in 1788 restricted the name A. Belladonna to the Cape Belladonna, excluding the literature. The same treatment was adopted independently by the younger Linnaeus in MS. 1781-83, though his work was not published until 1789. This treatment has been accepted without question for the past hundred years. We may therefore say that the name A. Belladonna should be retained for the Cape Belladonna, and the specimen in the Clifford Herbarium may well be accepted as the lectotype.


CybeRose Footnotes

1Miller (1768) gave the name A. Belladonna to the American plant; the Cape Belladonna he called A. Regina, after Linnaeus's A. Reginae. Linnaeus fils took his information from Hortus Nitidissimis, the relevant parts of which were completed before Seligmann's death in 1762, the year Linnaeus published Species Plantarum second edition. Peter Collinson (1746) and Dr. John Hill (1758) recognized that the plant of Sloane & Linnaeus was the Scarlet Belladonna from the West Indies.

2Miller (1731) described the West Indies, Italian and Portuguese Belladonnas as "very rare" in England. He noted that the Italian form was common around Florence, but also that the West Indies plant was "very common in the Barbadoes, St. Christopher's, and the other warm islands of the West Indies". Sir Hans Sloane had earlier written "It is planted along walks sides for ornament in gardens", which is to say that the West Indies Belladonna was a well known garden plant in Jamaica, a British territory.

3It should be remembered that Sloane had described the West Indies plant as polyanthos, and referred to the "several flowers". Formerly the English word "several" meant "separate". When it referred to a quantity, it meant "more than two". It would have been reasonable for Sloane to translate "several flowered" as "polyanthos", and for Linnaeus to translate this as "multiflora". Hermann mentioned "three, four or more flowers", and Du Tertre claimed that they produced five or six. Furthermore, John Parkinson (Paradisi) established the pattern of distinguishing Narcissus as monanthos, "one flower, or two at the most"; and polyanthos, "many flowers".

4Merian's plate, cited under Amaryllis Belladonna, has two open flowers and two buds. If the four-flowered Blank Specimen can be called "multiflora", there can be no objection to applying the same description to a four-flowered Amaryllis Belladonna.

5In Mantissa Plantarum Linnaeus added "Hemerocallidis"—like a daylily—to his description of Amaryllis Belladonna. In the 17th century, Rochefort and Hermann also commented on its resemblance to the Daylily.

6Miller (1731) discussed importing the Red Lily from the West Indies "in a box, dry" and described the cultural requirements. He noted that the Portuguese Belladonna Lily also had to be imported, because it did not survive long in English gardens.

7Miller (1759) wrote that the Zeylon Lily "is also tender, and must be treated in the same manner as the sixth (West Indies Belladonna); this is more common in the gardens in Holland than in this country; and as it is a plant which increases but slowly, will not be very common here." Therefore, we may not assume that the tenderer Scarlet Belladonna was more likely to have disappeared from the Leyden garden than the short-lived Cape plant.

8In 1760 Miller published Figures... with the plates reversed. The Scarlet Belladonna (t. 23) was printed with the Hort. Cliff. phrase name, as well as Sloane's. The Pink Belladonna (t. 24) was printed with Hermann's "Lilium Americanum ..." and Douglas's "Lilium Reginae". It was this edition that Linnaeus cited, which explains why he gave the name Amaryllis reginae to the Cape plant, and cited Hermann's phrase-name in connection with it. George Ehret had previously (1744) painted the Pink Belladonna with the name "Lilio-narcissus Americanus Belladona dictus par. bat." which is Plukenet's phrase-name, not that of Par. Bat. Sealy failed to mention that t. 23 (marginibus reflexis) is not the plant mentioned in the text of Plate XXIII (marginibus undulatis). Linnaeus could not read English, so he took his information from the plates and Latin phrases.

9Miller did not claim that he got his information from Sloane, only that the Belladonna lily was reported to have been collected by Sloane. Sloane described the hollow scape and wide-open flowers, which Sealy has chosen to ignore. Sloane cited: Lilium Americanum puniceo flora Belladonna dictum of Hermann, Lys rouge des Isles of Du Tertre, Lys des Antilles pareils a nos Lys jaunes, ou orangers of Rochefort, An Lilia rubra of Laet, and Red Lilly of Lignon. Rochefort compared the Red Lily he saw in Tobago to the orange Daylily. DuTertre also called the flowers "orange". All of these references, in addition to Miller (1731, 1741) show that Sloane was not describing an African Belladonna. The fact that Sloane's description was misapplied after the publication of Hortus Cliffortianus removes the need to denounce Linnaeus as a sloppy, uncritical, half-blind hack.

10Linnaeus cited the plates in Miller's Figures, but not the English descriptions. Linnaeus did not read English. In fact, Linnaeus cited the "Mexican Lily" under Amaryllis Belladonna and the "Belladonna lily" under A. reginae. Due to Miller's confusing text, and the reversal of the plates, Hermann's Belladonna was mistakenly included under A. reginae. Miller (1768), accepted Linnaeus's names, and cited his own plates as published. He returned Hermann's Belladonna to Amaryllis Belladonna.

11Sealy cleverly buried the phrase "Petala 3 superiora basi ciliata" in a long and tedious paragraph. It describes the beard pads of the American Belladonna, which are not found in the Cape plant. Dr. John Hill published an exquisitely detailed engraving of this feature in 1758, which was cited by Linnaeus in the appendix of Species Plantarum 2nd. (1762) and Mantissa Plantarum (1767 & 1771). The description Sealy mentioned was copied, with minor modifications, in the Mantissa.

12The Seligmann plate is titled "Lilio-narcissus", not "Amaryllis", and was copied from Ehret's 1744 painting mislabeled "Lilio-narcissus Americanus Belladona dictus par. bat." The text was not written by Trew, but by the classical scholar and linguist C. G. von Murr who borrowed freely from Miller's Dictionary.

13Amaryllis barbata is allied to A. Belladonna (Linn non Herb.) but is distinguished by several characters. It has white flowers, blooms in Summer rather than Spring, has much larger beard pads, etc. Alm had good reason to doubt whether this plant was identical to Merian's Lilium rubrum.

14Linnaeus stipulated that his son Carl was not to have the herbarium "as he never helped me in botany and has no love for it". After visiting Kew, Lin. fil. was sent home again, and bought his father's herbarium with borrowed money. The herbarium had been damaged by mildew and vermin, and Carl Jr. reportedly discarded badly damaged specimens. He died in 1783, just four years after his father.

15L'Héritier did not restrict Amaryllis Belladonna to the Cape Belladonna, nor did he suggest that he was using the names given by Linnaeus. In fact, he clearly indicated that he was not using Linnaeus's names by not appending "L." as he did for "Amaryllis formosissima L." Nor did he cite Species Plantarum or Miller's Dictionary (1768) for A. Belladonna and A. reginae as he did for some other species. In a letter to Dryander, L'Héritier wrote "You quarrel with me pitilessly over the name-changes. It is perhaps a bit too vague to accept always the first name that was printed. One runs the risk of adopting very bad ones, and this is of importance when one recognizes the possibility of giving a good one. (...) In general what should the trivial name be in order to be the best possible one? It would be necessary for it to distinguish the species from the others of the same genus, or at least to be applicable only to a single species."

As for Amaryllis Belladonna, L'Héritier included several species, including a Cybistetes and Aldini's Lilionarcissus rubeus indicus. He must have borrowed the latter misidentification from Plukenet because he also copied the misspelled "rubens". By reversing the names, L'Héritier transferred the ambiguity in Amaryllis reginae L. over to A. Belladonna L'Hérit.

See also: Refutation of Sealy