Typification of Linnaean names in Liliaceae

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Peruzzi & Jarvis • Linnaean names in Liliaceae

Typification of Linnaean names in Liliaceae Lorenzo Peruzzi1 & Charlie E. Jarvis2 1

Department of Biology, Unit of General and Systematic Botany, University of Pisa, Via Luca Ghini 5, 56126, Pisa, Italy. [email protected] (author for correspondence) 2 Department of Botany, Natural History Museum, Cromwell Road, London SW7 5BD, U.K. The typification of eleven Linnaean names of taxa belonging to the current circumscription of Liliaceae is discussed. Lectotypes are designated from the Linnaean herbarium at LINN (Lilium bulbiferum, L. candidum, L. martagon), the Burser herbarium at UPS (Bulbocodium serotinum, Fritillaria pyrenaica, Lilium bulbiferum), the Van Royen Herbarium at L (Fritillaria persica, Lilium chalcedonicum, Uvularia amplexifolia), the Clifford Herbarium at BM (Lilium pomponium) and from the Sloane herbarium in BM-SL (Bulbocodium autumnale). A specimen in BM is designated as the neotype of Lilium camschatcense. Each type choice maintains the present usage of the corresponding Linnaean name.

KEYWORDS: Bulbocodium, Fritillaria, Gagea, Lilium, Linnaean names, nomenclature, Streptopus, typification, Uvularia

INTRODUCTION The former broad concept of the family Liliaceae sensu lato (e.g., Cronquist, 1981) has, through the work of authors such as Dahlgren & al. (1985), Fay & Chase (2000), the Angiosperm Phylogeny Group (2003), Rønsted & al. (2005), Tan & al. (2005), Fay & al. (2006), Leitch & al. (2007) and Peruzzi & al. (2008, 2009), become more narrowly circumscribed in recent years. In the treatment of Peruzzi & al. (2009), which we follow here, the family now contains only 15 genera: Amana Honda, Calochortus Pursh, Cardiocrinum Endl. & Lindley, Clintonia Raf., Erythronium L., Fritillaria L., Gagea Salisb. (= Lloydia Salisb. ex Rchb.), Lilium L. (= Nomocharis Franchet), Medeola L., Notholirion Wallich ex Boiss., Prosartes D. Don, Scoliopus Torrey, Streptopus Michaux, Tricyrtis Wallich and Tulipa L. Approximately 850 species are included within the circumscription of Liliaceae sensu stricto, and 23 species names applicable to the group were established by Carl Linnaeus. Twelve of these names are already typified: Anthericum graecum L. (≡ Gagea graeca (L.) Irmisch), Fritillaria imperialis L., F. meleagris L., Lilium canadense L., L. philadelphicum L., L. superbum L., Medeola virginiana L., Ornithogalum luteum L. (≡ Gagea lutea (L.) Ker Gawl.), O. minimum L. (≡ Gagea minima (L.) Ker Gawl.), O. uniflorum L. (≡ Tulipa uniflora (L.) Besser ex Baker), Tulipa gesneriana L. and T. sylvestris L. (cf. Jarvis, 2007 and literature cited therein). The remaining eleven Linnaean names involved are, as far as we are aware, not yet typified and are therefore included in this study. Ten lectotypes and one neotype are designated to fix the application of these names according to current usage.

TYPIFICATION OF THE NAMES Bulbocodium autumnale L., Fl. Angl.: 14. 1754 ≡ Gagea serotina (L.) Ker Gawl. in J. Sci. Arts (London) 1: 180. 1816 ≡ Lloydia serotina (L.) Rchb., Fl. Germ. Excurs.: 102. 1830 – Lectotype (designated here): United Kingdom. Wales, Snowdonia, Glyder “Trigvulcaugh”, June 1700, R. Richardson s.n., Herb. Sloane 152: 156 (BM-SL!). [Treatment by Ray, 1724: 374:]

Binomials published in the Linnaean dissertation Flora Anglica (Linnaeus, 1754) are accompanied only by a number (in this case, “374.1”) which links the name to a species account in John Ray’s Synopsis Methodica Stirpium Britannicarum (Ray, 1724: 374, reproduced above; see also Stearn, 1973: 64–65). The binomial is therefore validated by Ray’s description and, under Art. 7.7 (see Ex. 8) of the Code, is to be typified from the context of Ray’s account rather than by using material in Linnaeus’s 1359


herbaria. Ray’s account is accompanied by an illustration (tab. XVII, fig. 1) and his text also contains reference to specimens collected in North Wales by Edward Llwyd (1660–1709) and Richard Richardson (1663–1741), now to be found in the Sloane Herbarium (HS 113: 224; 152: 156, BM-SL). We select here a complete and well-preserved specimen (HS 152: 156, BM-SL), collected by Richardson near Mt. Snowdon, as the lectotype of B. autumnale. The specimen, accompanied by Richardson’s handwritten label (transcribed by Dandy, 1958: 195), was originally in the herbarium of James Petiver (1658–1718). The type and the rest of the traced original material all correspond with the current concept of Gagea serotina (≡ Lloydia serotina). Bulbocodium serotinum L., Sp. Pl.: 294. 1753 ≡ Anthericum serotinum (L.) L., Sp. Pl., ed. 2: 444. 1762 ≡ Phalangium serotinum (L.) Poir. in Lamarck, Encycl. 5: 241. 1804 ≡ Gagea serotina (L.) Ker Gawl. in J. Sci. Arts (London) 1: 180. 1816 ≡ Rhabdocrinum serotinum (L.) Rchb., Consp. Regn. Veg.: 65. 1828 ≡ Lloydia serotina (L.) Rchb., Fl. Germ. Excurs.: 102. 1830 ≡ Cronyxium serotinum (L.) Raf., Fl. Tellur. 2: 28. 1837 – Lectotype (designated here): “In Vallesia in altissimo monte prope thermas Piperinas Helvetiorum: et in Taurero Rastadiensi, Burser Herbarium (III: 41, UPS!). [Protologue:]

Linnaeus took his diagnosis from van Royen (1740), and added a synonym from Ray (1724), and a second one from Bauhin (1620, 1623) and Rudbeck (1701). Adriaan van Royen’s Herbarium is conserved at L and was studied by Linnaeus. We traced there a herbarium sheet (No. 908.106-1769) bearing four individuals which are original material. Although the reference to Ray (1724) is identical to that which validates the later B. autumnale (1754, see above), only Ray’s figure (and not the specimens he cited) is original material for B. serotinum because Art. 7.7 does not apply in this case. Bauhin’s polynomial is linked with original material studied by Linnaeus in the Burser Herbarium (III: 41, UPS) and the plate depicted by Rudbeck (1701: 64, “IX Pseudonarcissus gramineo folio”) is also original material. Although the Linnaean Herbarium (LINN) contains two sheets (432.1 and 432.2; numbering follows Savage, 1945) 1360

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bearing material referable to this species, neither can be considered as original material since neither is annotated by Linnaeus and both carry only later annotations by James Edward Smith (1759–1828) indicating some doubt as to their identity: “Anthericum serotinum?”. Sheet 432.2 also carries a symbol normally linked with Siberia with Johann Georg Gmelin (1709–1755) as collector (cf. Stearn, 1957; Jarvis, 2007), and “Sibiria” is not mentioned in the protologue. We select here the well-preserved material in the Burser Herbarium (III: 41, UPS) as the lectotype of B. serotinum. The lectotype and the rest of the traced original material correspond with the current concept of Gagea serotina (= Lloydia serotina, e.g., Heywood, 1980). Fritillaria persica L., Sp. Pl.: 304. 1753 – Lectotype (designated here): Cultivated material, Herb. A. van Royen, No. 913.62-380 (L!). [Protologue:]

Linnaeus took his diagnosis from his own account in Hortus Upsaliensis (Linnaeus, 1748), adding synonyms from his Hortus Cliffortianus (Linnaeus, 1738) and also from van Royen (1740), Bauhin (1623) and Clusius (1601). There is no material associated with this name in any of the “general” Linnaean herbaria (LINN, S, etc.), and although there is in the Clifford Herbarium (sheet No. BM000558491, BM) a specimen identifiable as this species, it is not associated with the relevant entry there and cannot be considered as original material for the name. In the van Royen Herbarium there is a good herbarium specimen (No. 913.62-380, L) which can be considered as original material. There is another in the Burser Herbarium at UPS (“Florentiae in horto Magni Ducis”; III: 121a) but that specimen has been almost completely destroyed. There are also good figures in Clusius (1601: 130, “Lilium sufiani flor.” and “Lilium sufiani cum radice) which are original material too. We select here the complete and well-preserved specimen in the van Royen Herbarium as the lectotype of F. persica. The lectotype, and all the rest of the traced original material, correspond with the current concept of this species (e.g., Rix, 1980; Pignatti, 1982). Fritillaria pyrenaica L., Sp. Pl.: 304. 1753 – Lectotype (designated here): “Basileae in horto Eckenstenii ”, Herb. Burser III: 62 (UPS!).

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[Protologue:]

In the Linnaean Herbarium at LINN there is a sheet (No. 421.2) marked “pyrenaica. 4 ” by Linnaeus and therefore acceptable as original material. However, it is not identifiable as F. pyrenaica as currently understood. In S, there is a specimen (No. 139.19) marked as Fritillaria pyrenaica but it is annotated solely by Lars Montin (1723–1785) and is a post-1753 addition. Linnaeus also gave earlier synonyms from his own Hortus Upsaliensis (Linnaeus, 1748: 81, as “Hort. cliff. ” but clearly in error for “Hort. ups.” as already noted by López González, 1986), and from Bauhin (1623) and Clusius (1601, 1611: 10 – a mistake for p. 20). In the Burser Herbarium at UPS there is a sheet (III: 62) associated with this name that López González (1986) considered ineligible as lectotype, arguing that it conflicted with Linnaeus’s diagnosis “foliis infimis oppositis ”. The specimen indeed has only subopposite basal leaves, but it also has a pair of apparently opposite leaves in the median portion of the stem. In our opinion, there is not a definite conflict with the protologue, also considering that the Clusius (1601: CCLVI–CCLVII) account cited by Linnaeus also gave a full and detailed description that is compatible with the Burser material. We select here the complete and well-preserved Burser specimen to serve as the lectotype of F. pyrenaica. The lectotype corresponds with the traditional concept of this endemic of northern Spain and southern France. Fritillaria pyrenaica has been treated as a nomen confusum by some recent authors (e.g., Fernandez-Arias Gonzalez & Devesa Alcaraz, 1990) and informally rejected in favour of F. nervosa Willd. However, no formal proposal for the rejection of the Linnaean name appears to have been made and we favour retaining the latter. Lilium bulbiferum L., Sp. Pl.: 302. 1753 – Lectotype (designated here): Herb. Linnaeus, sheet No. 420.2 (LINN!). [Protologue:]


Linnaeus cited the diagnosis as coming from his Hortus Cliffortianus (Linnaeus, 1738), also cited via his Hortus Upsaliensis (Linnaeus, 1748), van Royen (1740) and Gmelin (1747). He also added eight polynomials from Bauhin (1623) as synonyms, of which seven were noted as unnamed varieties (as indicated by an appended Greek letter). In the Linnaean Herbarium at LINN there is a sheet (No. 420.2) marked as “bulbiferum. 4 ” by Linnaeus which, though original material and with a few bulbils, has badly damaged flowers. In the Clifford Herbarium, there is one sheet identifiable as L. bulbiferum (No: BM000558496, a plant lacking bulbils) but it is not associated with any account in that book and cannot be considered original material. In the van Royen Herbarium, a specimen (No. 913.62-583, L) is original material and another, in the Burser Herbarium at UPS (III: 115, “Lilium purpureocroceum majus”, a plant lacking bulbils) is associated with the first of the Bauhin polynomials from Linnaeus’s protologue. Another Burser specimen (III: 116, “Lilium bulbiferum majus”, a plant bearing bulbils) is linked to the unnamed variety ε. We select here the material in LINN to serve as the nomenclatural type of L. bulbiferum. Although it has suffered some insect damage, it does bear bulbils and therefore corresponds with the traditional application of this name. Plants lacking bulbils (which are very common, for instance, in peninsular Italy) are currently called L. bulbiferum subsp. croceum (Chaix) Jan (e.g., Matthews, 1980; Pignatti, 1982; Conti & al., 2005). Lilium camschatcense L., Sp. Pl.: 303. 1753 ≡ Fritillaria camschatcensis (L.) Ker Gawl. in Bot. Mag. 30: ad t. 1216. 1809 – Neotype (designated here): Russia: “Siberia”, Rudolph s.n. (BM!). [Protologue:]

Linnaeus cited the diagnosis from his own account in Amoenitates Academicae (Linnaeus, 1751: 348) where there is a detailed description of this species. This account is itself a slightly modified reprinting of an original dissertation, Plantae Rariores Camschatcenses (Linnaeus, 1750), which described a collection of herbarium specimens, ostensibly from Kamchatka, that had been sent to Linnaeus on loan by Grigory Demidov (see Sokoloff & al., 2002: 163–169; Jarvis, 2007: 89 for further information). Although Linnaeus retained some duplicates, these appear not to have included a specimen of this species, and the original specimen was presumably returned to Demidov, 1361


and has subsequently been lost. Linnaeus also indicated “Canada” as a source of material of this species, and provided notes on the differences between the Kamchatka and North American material. “Canada” for Linnaeus was chiefly the area of eastern North America explored by his student Pehr Kalm. This extended “roughly from Philadelphia and New York northward, by way of Albany, to Montreal and Quebec and from Albany westward along the Mohawk River to Lake Ontario and Niagara Falls” (Stearn, 1957: 144). Although there is a Kalm specimen (420.7, LINN) in existence, it was annotated “philadelphicum” by Linnaeus, and is the lectotype (designated by Reveal in Jarvis, 2007: 632) of Lilium philadelphicum L. (1762) which Linnaeus, recognising that it was distinct, later segregated from L. camschatcense. The latter species is present in North America only to the west of the Rocky Mountains (see Ness, 2002: 168), so Kalm could not have encountered this species so far to the east. Matsuura (1935), studying Japanese populations, showed that slender plants from high mountain habitats are diploid, while robust plants from lowlands are triploid. Subsequently, Matsuura & Toyokuni (1963) recognised the diploid taxon as Fritillaria camschatcense subsp. alpina Matsuuri & Toyok., with susbsp. camschatcense applied to the lowland triploid populations. Further information on the distribution and synonymy of this species is provided by Turrill (1949) and Sealy (1980). We have been unable to locate any original material for this name which therefore, as already noted by Matsuura & Toyokuni (1963), requires a neotype. As material from Kamchatka corresponds with Linnaeus’s original description, and is apparently triploid, we designate a neotype from this region, corresponding with the typical subspecies sensu Matsuura & Toyokuni.

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there is a sheet (No. 420.1) marked “candidum 1” by Linnaeus, which has badly damaged flowers, while in S there is only a post-1753 specimen (No. 139.13) annotated solely by Lars Montin. There is no relevant material in the Clifford Herbarium (BM). In the van Royen Herbarium (No. 913.62-563, L) there is a relevant specimen but it is, unfortunately, sterile. Further original material is in the Burser Herbarium (Lilium album, flore erecto et vulgare, III: 113, UPS) but that specimen is also badly damaged. Finally, there is a cited figure from Dodoëns (1583: 197), and a further Burser specimen (III: 114) associated with variety β. Dasgupta & Deb (1984: 488) indicated 421.1 (LINN) as the type, but this is material linked with Fritillaria and cannot be original material for L. candidum. Although 420.1 (LINN) has been damaged by insects, we nevertheless prefer to select it as the lectotype rather than Dodoëns’s illustration. We have no doubts as to the taxonomic identity of the type specimen, which will also serve as the generitype of Lilium. The lectotype and the traced original material correspond with the current concept of this species (e.g., Matthews, 1980), which is also one of the most ancient ornamental plants of the Western World (Tamura, 1998). Lilium chalcedonicum L., Sp. Pl.: 302. 1753 – Lectotype (designated here): Herb. A. van Royen (sheet No. 913.62-556, L!). [Protologue:]

Lilium candidum L., Sp. Pl.: 302. 1753 – Lectotype (designated here): Herb. Linnaeus, sheet No. 420.1 (LINN!). [Protologue:]

Linnaeus cited his diagnosis from his own Hortus Cliffortianus (Linnaeus, 1738), also cited via his Hortus Upsaliensis (1748) and Materia Medica (Linnaeus, 1749), and via van Royen (1740). Synonyms from Bauhin (1623) and Dodoëns (1583) were also given, as was a polynomial from Bauhin (1623), which Linnaeus associated with an unnamed variety β. In the Linnaean Herbarium (LINN) 1362

In the Linnaean Herbarium (LINN), sheet 420.4 is marked “4” (the number of L. chalcedonicum in Species Plantarum) by Linnaeus with “chalcedonicum” added later by Smith. It is original material for the name, but it has suffered some insect damage. Another sheet in LINN (No. 420.5) is marked “chalcedonicum” by Linnaeus, but also with the number “5” (the number of L. martagon L. in Species Plantarum). The sheet also carries a symbol indicating the western edge of Asia and, on the verso, Traugott Gerber (1710–1743) indicated as the collector, along with “Habitat in campis feris inter Jelez & Woroniz etiam circa Tawrow” written by Linnaeus. As the specimen is identifiable as L. martagon, it seems that there

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was a labelling error between number and epithet, and that this sheet is original material for L. martagon, not L. chalcedonicum. In the Clifford Herbarium (BM) are three minimally annotated sheets (BM-000558493, BM-000558494, BM000558495) that have been linked with Lilium 4 in the account in Hortus Cliffortianus but because, in Species Plantarum, Linnaeus also cited the same polynomial as a synonym of L. pomponium, the Clifford specimens could potentially be original material for either name. The diagnoses of these two names in 1753 differ subtly but significantly from one another with “[foliis sparsis] subulatis” used for L. pomponium, and “[foliis sparsis] lanceolatis” used for L. chalcedonicum. However, all three Clifford sheets clearly represent the plant currently called L. pomponium. In the van Royen Herbarium, there is one relevant herbarium specimen (No. 913.62-556, L) which appears to be original material and is identifiable as Lilium chalcedonicum. The figure depicted in Clusius (1601: 131; “Lilium rubrum sive miniatum byzantinum”) is also original material. In the Burser Herbarium there is a specimen (Lilium purpuro-sanguineum, flore reflexo, III: 118, UPS) linked to the unnamed variety γ, as is the figure in Lobelius (1581: 169; “Hemerocallis chalcedonica”). The extant specimens have all suffered some degree of insect damage, but we select that in the van Royen Herbarium to serve as the nomenclatural type of L. chalcedonicum. It and the rest of the original material (excluding that associated with var. γ) that we have traced correspond with the current concept of this species (e.g., that of Matthews, 1980), a taxon endemic to S Albania and Greece, and clearly distinguished from other Turk’s cap lilies in having leaves changing abruptly in aspect from patent below to appressed above. Linnaeus’s choice of epithet suggests he believed the plant came from northwestern Anatolia (ancient village of Chalcedon), and his synonymy indicates plants coming from the Istanbul area. It is likely, however, that this beautiful species was already cultivated in that area at that time. Linnaeus’s var. γ probably represents a red-flowered form of a taxon within the L. carniolicum Bernh. ex Koch group, such as Lilium ponticum C. Koch or L. artinense Miscz. (both of which are endemic to northern Anatolia, cf. Davis & Henderson, 1984). It is interesting that the illustration in Clusius is a copy of a figure originally published by Lobelius (1581: 169; “Hemerocallis byzantina altera”) together with the above-mentioned figure “Hemerocallis chalcedonica” (which applies to another taxon in the L. carniolicum group). Lilium martagon L., Sp. Pl.: 303. 1753 – Lectotype (designated here): Gerber s.n., Herb. Linnaeus, sheet No. 420.5 (LINN!).


[Protologue:]

Linnaeus cited his diagnosis from his own Hortus Cliffortianus (1738), also cited via his Hortus Upsaliensis (1748), Gmelin (1747) and van Royen (1740). Synonyms came from Bauhin (1623) and Dodoëns (1583), with an unnamed variety β based on a further Bauhin polynomial. As discussed above under L. chalcedonicum, sheet No. 420.5 (LINN) appears to be original material for L. martagon, unlike a specimen in the Linnaean Herbarium in S (No. 139.15), which is a post-1753 addition annotated solely by Lars Montin. There is no relevant material in the Clifford Herbarium (BM) but there is original material in the van Royen Herbarium (No. 913.62-570, L), comprising only an inflorescence. In the Burser Herbarium (III: 117, UPS) there is material only doubtfully associated with either of the polynomials cited by Linnaeus, though Linnaeus did independently confirm his view of the identity of the specimen (see Savage, 1937). The figure depicted in Dodoëns (1583: 201) is also original material. We select the specimen in the Linnaean herbarium (LINN) to serve as the lectotype of Lilium martagon. The lectotype and the rest of the traced original material correspond with the current concept of this species (e.g., Matthews, 1980). Lilium pomponium L., Sp. Pl.: 302. 1753 – Lectotype (designated here): Clifford Herbarium: 120, Lilium 4C (BM-000558495). [Protologue:]

Linnaeus cited the diagnosis from his own Hortus Cliffortianus (1738), also cited via his Hortus Upsaliensis (1748) and van Royen (1740). Synonyms from Gmelin (1747), Bauhin (1623) and Clusius (1601) are cited, along with two polynomials from Bauhin (1623) associated with unnamed varieties β and γ, respectively. In the Linnaean Herbarium (No. 420.3, LINN) there is a sheet marked “Mart. pomponium” by Linnaeus but 1363


it lacks the relevant Species Plantarum number (i.e., “3”) and is evidently a post-1753 addition to the collection. As discussed above under L. chalcedonicum, there are three sheets (BM-000558493, BM-000558494, BM000558495) in the Clifford Herbarium (BM) linked with L. pomponium. The figure depicted in Clusius (1601: 133; “Lilium rubrum praecox”) is also original material. The unnamed variety β is represented by cultivated material in the Burser Herbarium at UPS (Lilium miniatum odorum angustifolium, III: 119, UPS) which does not appear to be identifiable as L. pomponium (see also Juel, 1923: 41). We select here one of the Clifford Herbarium sheets as the lectotype of Lilium pomponium. It and the rest of the traced original material (apart from that associated with var. β) correspond with the current concept of this species, endemic to the Maritime Alps (Matthews, 1980). Linnaeus’s indication of the Pyrenees and Siberia as part of the distribution of this species is erroneous. Uvularia amplexifolia L., Sp. Pl.: 304. 1753 ≡ Streptopus amplexifolius (L.) DC. in Candolle & Lamarck, Fl. Franç., ed. 3, 3: 174. 1805 – Lectotype (designated here): Cultivated material, Herb. A. van Royen, No. 913.62-465 (L!). [Protologue:]

Linnaeus coined a new diagnosis and also cited synonyms from van Royen (1740) and Barrelier (1714) with a third from Bauhin (1623), also cited via Bauhin & Cherler (1651), Morison (1699) and Ray (1724), a fourth from Clusius (1601) and a fifth from Matthiolus (1586). There is no relevant material in any of the general Linnaean herbaria, but there is original material (No. 913.62-465, L) in the van Royen Herbarium, and in the Burser Herbarium (XVII: 63, UPS). Cited illustrations are present in Barrelier (1714: 58, t. 720), Bauhin & Cherler (1651: 530), Morison (1699: 537, s. 13, t. 4, f. 11), Ray (1724: 665), Clusius (1601: 276) and Matthiolus (1586: 936). We select the well-preserved, fruiting specimen from Adriaan van Royen as the lectotype of Uvularia amplexifolia. The lectotype and all the other traced original material correspond with the current concept of Streptopus amplexifolius, the only species of this genus occurring in Europe (Valentine, 1980). 1364

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ACKNOWLEDGEMENTS We are most grateful for helpful comments made by Nicholas Turland (MO) on an earlier version of this manuscript and to the Nomenclature Editor, John McNeill, for considerably improving our original treatment of Lilium camschatcense.

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