Introduction
Eriocaulon L. is a large genus of c.400 species distributed widely in the tropics and subtropics (Stützel, Reference Stützel and Kubitzki1998; Mabberley, Reference Mabberley2008). It is the only genus of Eriocaulaceae that occurs in Southeast Asia. The plants are usually small, rosulate herbs that may be annual or perennial. Most are found in open areas that are permanently or seasonally damp.
Lecomte (Reference Lecomte1912) recognised 27 species of Eriocaulon from Cambodia, Laos and Vietnam. More recently, Zhang's monograph of the species of East Asia (1999) recognised 28 species and 1 variety of Eriocaulon in Thailand and 10 species in Cambodia, Laos and Vietnam. Neither of these works addressed questions of typification in detail. During preparation of revisions of Eriocaulon for the Flora of Thailand and the Flora of Cambodia, Laos and Vietnam, a number of names have been treated for which lectotypes are required.
Five names requiring typification were coined by Lecomte in J. Bot. (Morot), sér. 2, 1 (Reference Lecomte1908a,b), namely, Eriocaulon alatum Lecomte (Reference Lecomte1908b, p. 104), Eriocaulon lanigerum Lecomte (Reference Lecomte1908b, p. 109), Eriocaulon nautiliforme Lecomte (Reference Lecomte1908a, p. 89), Eriocaulon nigrum Lecomte (Reference Lecomte1908b, p. 107) and Eriocaulon ubonense Lecomte (Reference Lecomte1908b, p. 109). Lecomte (Reference Lecomte1908a,Reference Lecomteb) published two papers on Eriocaulon in série 2, volume 1, of the Journal de Botanique.
The first paper, Eriocaulacées de Chine et d'Indo-Chine de l'Herbier du Muséum, appeared on pp. 86–94 in part 4 of the volume in April 1908, and the second, Espèces nouvelles d'Eriocaulon de l'Indo-Chine, came out on pp. 101–109 in part 5, dated May 1908. On p. 89 of the earlier paper, there is a key to species that includes the names Eriocaulon nautiliforme, E. nigrum and E. ubonense. Of these, only Eriocaulon nautiliforme can be considered to be validly published, because the characters given in the key clearly and uniquely separate this species from all others (Art. 32.1(d) in McNeill et al., Reference McNeill, Barrie, Buck, Demoulin, Greuter, Hawksworth, Herendeen, Knapp, Marhold, Prado, Prud'homme van Reine, Smith, Wiersema and Turland2012). Eriocaulon nigrum, by contrast, comes out in two places in the key, each time in a group of species, and is not separated from these others. Eriocaulon ubonense likewise keys out with other species. This key also includes a reference to Eriocaulon australe, but in Lecomte's second paper of 1908, he described this taxon as E. lanigerum without mentioning that he had previously called it E. australe. Although Eriocaulon nautiliforme is validly published on p. 89, no type is cited.
Lecomte clearly intended to publish his new species in the second paper. The valid publication of Eriocaulon nautiliforme in the first paper is incidental although correct. Syntypes are cited in the second paper, on p. 106. The place of valid publication of Eriocaulon alatum, E. lanigerum, E. nigrum and E. ubonense is the second paper, where syntypes are cited.
Two other names, Eriocaulon hamiltonianium Mart. in Wall., Pl. Asiat. Rar. 3: 29 (Reference Martius and Wallich1832) and Eriocaulon infirmum Steud., Syn. Pl. Glumac. 2: 271 (Reference Steudel1855), involve Karl Friedrich Philipp von Martius and Nathanial Wallich. Wallich distributed specimens collected by a number of workers on behalf of the East India Company; many collections are known from a number of duplicates that are now located in different herbaria. The ‘top’ set of the East India Company Herbarium was kept in London and is now at the Royal Botanic Gardens, Kew, where it is held as a separate collection (K-W).
Wallich's Numerical List (Reference Wallich1829–Reference Wallich1849) was originally designed to be cut up and used as specimen labels for the specimens he distributed (Wallich Catalogue Online, 2008–). It contains 9148 entries, usually ‘species’, and gives the collecting details of 20,897 ‘collections’ (Wallich Catalogue Online, 2008–). It includes material collected by Francis Hamilton (= Francis Buchanan-Hamilton), although there are unlikely to be many duplicates of material collected by Buchanan-Hamilton, because Wallich noted in his Numerical List (Reference Wallich1829–Reference Wallich1849) that the Buchanan-Hamilton herbarium had very few duplicates. Buchanan-Hamilton's own herbarium was given to the University of Edinburgh on his death and is now at the Royal Botanic Garden Edinburgh (E).
Von Martius was likely responsible for deciding on the grouping of Eriocaulaceae collections in Wallich's Numerical List (Reference Wallich1829–Reference Wallich1849) into species and for providing their names, so he was sent the material before the collection was numbered. Therefore, relevant material from his personal herbarium is embossed ‘HERBARIUM C. F. Ph. Martii’ but does not include the Wallich Numerical List number. Martius's private herbarium was sold on his death, and all his Wallich specimens went to Meise, Belgium (BR; H.-J. Esser, pers. comm. and M. F. Watson, pers. comm.).
Specimens were studied at the Natural History Museum, London (BM), including the Hermann collection; the Botanic Garden Meise (BR); the Royal Botanic Gardens, Kew (K), including the Wallich Herbarium (K-W); the Linnean Society (LINN); the Muséum National d'Histoire Naturelle, Paris (P); and the herbarium of the Institute of Tropical Biology, Ho Chi Minh City (VNM). Unless otherwise indicated, all specimens have been seen. Our methods follow the International Code of Nomenclature for algae, fungi, and plants (Melbourne Code) (McNeill et al., Reference McNeill, Barrie, Buck, Demoulin, Greuter, Hawksworth, Herendeen, Knapp, Marhold, Prado, Prud'homme van Reine, Smith, Wiersema and Turland2012).
Lectotypification
Eriocaulon alatum Lecomte, J. Bot. (Morot) sér. 2, 1: 104 (Reference Lecomte1908b) & Fl. Indo-Chine 7(1): 17 (Reference Lecomte1912); Merr., Fl. Manila: 136 (Reference Lecomte1912); P.H. Hộ, Ill. Fl. Vietnam 3(1): 480 (1993). – Type: COCHINCHINE. L. Pierre s.n. (lectotype, P [P01721442], designated here; isolectotypes K [K000960552]; P [P01721443, P01721444 (pro parte), P01721445]).
Eriocaulon alatum is a synonym of Eriocaulon zollingerianum Körn. (Royen, Reference Royen1959), but no lectotype of E. alatum has been designated. Lecomte cited two collections in the protologue, namely Thorel s.n. from Laos (Bassac) and Pierre s.n. from southern Vietnam (Cochinchine). We have located specimens of Pierre s.n. at P and K, but we have found no specimens of Thorel s.n. from Bassac. The specimen of Pierre s.n. at P [P01721442] is the best preserved of the materials seen, is the most appropriate choice as lectotype and is therefore chosen as the lectotype. There are four isolectotypes, one at K [K000960552, which was cited as a syntype by Zhang (Reference Zhang and Kück1999: 187)], and three at P [P01721443, P01721444 & P01721445]. Pierre s.n. [P01721444] is a mixed collection, the lower right-hand specimen being Eriocaulon nepalense Prescott ex Bong. and the rest of the material isolectotypes of E. alatum. We note that Zhang (Reference Zhang and Kück1999: 187) also cited Pierre s.n. (G) as a syntype of Eriocaulon alatum. We have only seen images of two such sheets annotated by Zhang [G00191983 & G00191984] and cannot confirm that they are part of the same gathering as the lectotype.
Eriocaulon hamiltonianum Mart. in Wall., Pl. Asiat. Rar. 3: 29 (Reference Martius and Wallich1832); Kunth, Enum. Pl. 3: 552 (1841); Steud., Syn. Pl. Glumac. 2: 270 (Reference Steudel1855); Körn., Linnaea 27: 679 (1856); Ruhland in Engl., Pflanzenr. IV, 30 (Heft 13): 107 (Reference Ruhland and Engler1903); Z.X.Zhang in U.Kück, Diss. Bot. 313: 169–172 (Reference Zhang and Kück1999); R.Ansari & N.P.Balakr., Fam. Eriocaul. India: 122 (Reference Ansari and Balakrishnan2009). – Type: INDIA. Gualpara, F. Hamilton, s.n. (lectotype BR [BR000000918773], designated here; isolectotypes, E [E00027135]; K-W [K001122954]).
Eriocaulon hamiltonianum is based on a collection made by Buchanan-Hamilton from Gualpara [Goalpara], Assam, North India, appearing in Wallich's Numerical List under entry 6075 (p. 207). There are three potential lectotypes.
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• Martius's private herbarium in BR includes one specimen of Eriocaulon hamiltonianum. This specimen bears a label that reads, ‘Eriocaulon cinereum Herb. Hamilt. e Gualpara’ in Wallich's hand, and ‘Hamiltonianum Mart.’ in Martius's hand. The label is embossed ‘HERBARIUM C.F.Ph. Martii’ but does not include the Wallich Numerical List number 6075.
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• Both Martius and Wallich recorded that the Buchanan-Hamilton collection was given to the East India Company Herbarium under Buchanan-Hamilton's field name Eriocaulon cinereum. There is also one specimen in K-W, number 6075, bearing Buchanan-Hamilton's label ‘E. cinereum Gualpara 4th Nov. 1808’ at the top left, and ‘6075. E. hamiltonianum Mart. E. cinereum H. Ham. e Gualpara’ at the top right.
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• The other specimen in Buchanan-Hamilton's own herbarium at the Royal Botanic Garden Edinburgh (E [E00027135]) bears Buchanan-Hamilton's field ticket, which reads, ‘Eriocaulon cinereum, Gualpara, 4th Nov. 1808’, and a printed herbarium label with Buchanan-Hamilton's Bengal Survey Catalogue number ‘348’ and a transcription of the information from this Catalogue.
Because Martius did not specify which herbarium he was using, a lectotype must be selected from the three known elements that comprise the original material listed above. Martius would not have seen the specimen in Buchanan-Hamilton's herbarium, which cannot, therefore, be considered. As an ‘Assignee’, Martius may have been sent all the material of this collection in the East India Company by Wallich, and so he may have seen the material in K-W, returning most of it after identification and keeping one part for himself. Because the duplicate at BR is the only one bearing the name Eriocaulon hamiltonianum in Martius's hand, we choose this one.
Eriocaulon hookerianum Stapf, Trans. Linn. Soc. London, Bot. 4: 243 (Reference Stapf1894); Ruhland in Engl., Pflanzenr. IV, 30 (Heft 13): 83 (Reference Ruhland and Engler1903); Lecomte Fl. Indo-Chine 7(1): 10 (Reference Lecomte1912); P.H. Hộ, Ill. Fl. Vietnam 3(1): 392 (1993); Beaman & R.S. Beaman (Reference Beaman and Beaman1998: 137); Z.X.Zhang in U.Kück, Diss. Bot. 313: 143 (Reference Zhang and Kück1999); R.Ansari & N.P.Balakr., Fam. Eriocaul. India: 28 (Reference Ansari and Balakrishnan2009). – Type: BORNEO. Mount Kinabalu, Temburungo, 7700 ft, C.D. Haviland 1204, excluding attached line drawing made from Haviland 1153, individual at top right of sheet (lectotype K [K000291319], designated here).
Otto Stapf worked on Dr Haviland's collections from Sabah soon after his appointment to the herbarium at Kew (Anonymous, 1933). Stapf, in the protologue (1894), cited five collections from two locations under Eriocaulon hookerianum: Burbidge s.n., Low s.n. and Haviland 1153 from Maripari Spur, Sabah, and Haviland 1153 bis and 1204 from Temburungo, Sabah. Beaman & Beaman (Reference Beaman and Beaman1998) listed these syntypes as follows, “Type: Kemburongoh, 2300 m, Haviland 1153 bis (syntype K n.v.), 1204 (syntype K n.v.); Marai Parai, Low s.n. (syntype fragment L!); Marai Parai Spur, 1500–1700 m, Burbidge s.n. (syntype K n.v.) 1500–1700 m, Low s.n. (syntype K n.v.; syntype fragment L!). The Kew specimens apparently have been misplaced.” Beaman & Beaman (Reference Beaman and Beaman1998) use the modern place name spellings Kemburongoh and Marai Parai, whereas the original labels give Temburungo and Maripari, or Marie Parie. The collector Low is written Lowe on the original labels, but according to van Steenis (Reference Steenis1950), the collector was Hugh Low (1824–1905) and Lowe is a misspelling. We have seen all these collections at K except Haviland 1153 bis, but neither of the fragments at L cited by Beaman & Beaman (Reference Beaman and Beaman1998). Among the specimens now at K, which would have been seen by Stapf, Haviland 1204 clearly shows the characters of the species and is better preserved than the remaining specimens. Therefore, we select it as the most appropriate choice of lectotype. All the plants attached to this sheet are of Eriocaulon hookerianum, but in order to avoid any further doubt, we select one individual as the lectotype.
Ansari & Balakrishnan (Reference Ansari and Balakrishnan2009) treated Eriocaulon hookerianum, citing the type as follows: “Type: Haviland 1153, Maripari Spur, N. Borneo, 5000–5500 ft. (K)”. They also noted that, “E. hookerianum is originally a Bornean species. We could not examine the type.” There is no suggestion that Ansari & Balakrishnan intended to select a lectotype, nor did they use the phrase “designated here” or the equivalent, as demanded by the International Code of Nomenclature since 1 January 2001.
Eriocaulon infirmum Steudel (Reference Steudel1855: 271); P.Royen, Nova Guinea 10(1–2): 38 (Reference Royen1959).
≡ Eriocaulon gracile Mart. in Wall., Pl. Asiat. Rar. 3: 29 (Reference Martius and Wallich1832), non Eriocaulon gracile Bong., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 1: 634 (Reference Bongard1831). – Type: BURMA. Mountains of Prome, Irawaddi, 1826 [1827], N. Wallich s.n., Numerical List number 6079 (lectotype BR [BR0000008570581], designated here; isolectotype K [K000433038]; K-W [K001122959]).
Steudel (Reference Steudel1855) published the name Eriocaulon infirmum for Eriocaulon gracile Martius (Reference Martius and Wallich1832). Although Steudel did not say so explicitly, his name Eriocaulon infirmum is a replacement name, because Eriocaulon gracile Bongard (Reference Bongard1831) had already been published, based on a different type. Bongard's species is from Brazil and is now called Syngonanthus gracilis (Bong.) Ruhland.
Martius (Reference Martius and Wallich1832: 29) cited “Wallich 1827” as the type, but no specimen marked with this year of collection has been found by us. Probably, a typographic error occurred in publication. Wallich's Numerical List refers to this collection under entry 6079 and gives the year of collection as 1826. The specimen selected as the lectotype of Eriocaulon gracile is held at BR and bears a label, “Herbarium Martii” with the date 1828. Above this is a smaller label in Wallich's hand with the collection date 1826. The reason for preferring the specimen at BR over the one at K-W is given under Eriocaulon hamiltonianum above.
Eriocaulon lanigerum Lecomte, J. Bot. (Morot) sér. 2, 1: 109 (Reference Lecomte1908b); Fl. Indo-Chine 7(1): 8 (Reference Lecomte1912). – Type: COCHINCHINE. 1862, C. Thorel 1493 (lectotype P [P00224173], designated here; isolectotype P [P00745124]).
Lecomte cited two collections in the protologue, namely Thorel 1493 and Pierre s.n., both from Cochinchine. Of these, only the first can be found at P. There are two duplicates, P00224173 & P00745124; of these, P00224173 is the better preserved and is chosen as the lectotype. P00745124 is, therefore, an isolectotype. As mentioned above, Eriocaulon lanigerum was treated as Eriocaulon australe in Lecomte's first paper, but E. lanigerum is a very small annual plant with soft heads of pilose bracts whereas E. australe is a more robust perennial plant with hard heads of glabrous bracts. Note that Eriocaulon australe is now treated as Eriocaulon sexangulare var. australe (R.Br.) Praj. & J.Parn. (Prajaksood et al., Reference Prajaksood, Parnell and Chantaranothai2012).
Eriocaulon nautiliforme Lecomte, J. Bot. (Morot) sér. 2, 1: 89 (Reference Lecomte1908a); J. Bot. (Morot) sér. 2, 1: 105–106 (Reference Lecomte1908b); Fl. Indo-Chine 7(1): 7 (Reference Lecomte1912). – Type: CAMBODIA. Kampot, 6 December 1903, C. Geoffray 215 (lectotype P [P00224175], designated here).
Lecomte (Reference Lecomte1908a) referred to three collections under this name: Thorel 1593, collected from “Bassac, dans le Me-Kong”; Pierre s.n., from Vietnam, Chaudoc; and Geoffray 215, from fallow rice fields. We have located the following sheets of these collections:
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• Geoffray 215 [P00224175], collected at Kampot, Cambodia
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• Pierre s.n. [P00224176], collected at Chaudoc, Vietnam
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• Thorel 1593 [B 10 0086998], collected at Bassac, Laos [image seen at the Virtual Herbarium Berolinense (Botanischer Garten und Botanisches Museum Berlin-Dahlem, no date)]
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• Thorel 1593 [K, without barcode number], collected at Bassac, Laos
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• Thorel 1593 [P00224177], collected at Bassac, Laos
Of the three type collections seen, Geoffray 215 is the best preserved and is the most appropriate choice as lectotype.
Pierre s.n. [P00224176] is a syntype of Eriocaulon nautiliforme.
Thorel 1593 (B) is a syntype of Eriocaulon nautiliforme. Zhang (Reference Zhang and Kück1999) states, under the title Untersuchte Belege, “THOREL s.n. Laos: Bassac, 1893, Syntypus B”. This indicates that he knew of other type specimens but had not studied them and did not intend to select a lectotype. His use of the term syntype confirms this. A high-resolution image of this sheet may now be seen at the Virtual Herbarium Berolinense. Examination of this image reveals that it is a duplicate of Thorel 1593. The number 5 in late 19th century handwriting is very easy to mistake for an 8. We believe that Zhang misread the collection number, thinking that it might be a collection date.
There are four sheets of Thorel 1593 at P. [P00224183] is a syntype of Eriocaulon ubonense. Lecomte clearly recognised that Thorel 1593 was a mixed collection (see below). The sheets with barcodes P01729272 & P01729278 have been determined to Eriocaulon odoratum Dalzell (P. Souladeth, 19 May 2014) and have been annotated Thorel 1593a. The sheet with barcode P01729276 is a mixed collection and has been annotated Thorel 1593b (P. Souladeth, 19 May 2014). Most of the material on sheet P01729276 is Eriocaulon siamense Moldenke var. macrophyllum Praj. & J.Parn., whereas one scape is E. zollingerianum (P. Souladeth, 19 May 2014).
Finally, there is a duplicate of Thorel 1593 at K, without a barcode number. This is a syntype of Eriocaulon nautiliforme.
Eriocaulon nigrum Lecomte, J. Bot. (Morot) sér. 2, 1: 107 (Reference Lecomte1908b); Fl. Indo-Chine 7(1): 16 (Reference Lecomte1912). – Type: VIETNAM. Western Tonkin, “In arenosis agris Minh Chinh”, 16 December 1881, H. Bon 1136 (lectotype P [P00224179], designated here; isolectotype P [P00745125]).
Two specimens collected from Vietnam were cited under this name: Bon 1136 [P00224179 & P00745125], collected in 1881, and Balansa 243 [P00224178 & K000959944], collected in 1885. Bon 1136 clearly shows the characters of the species described in the protologue and is better preserved than Balansa 243. Therefore, Bon 1136 is the more appropriate choice as the lectotype; Balansa 243 remains a syntype.
Eriocaulon ubonense Lecomte, J. Bot. (Morot) sér. 2, 1: 109 (Reference Lecomte1908b); Fl. Indo-Chine 7(1): 11 (Reference Lecomte1912); P.H. Hộ, Ill. Fl. Vietnam 3(1): 398 (1993); Koyama, Philipp. J. Sci. 84(3): 373 (1955). – Type: CAMBODIA. October, 1874 L. Pierre s.n. (lectotype P [P00224182], designated here).
Lecomte (1908) cited two collections under this name: Thorel 1593, collected from Ubon Ratchathani, Thailand, and Pierre s.n., collected from Cambodia. The following specimens have been seen only at P; no duplicates have been found.
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• Pierre s.n. [P00224182]. The label states “[Cambodia 10/1874]” in a modern hand.
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• Thorel 1593 [P00224183]. The place name “Ubon” has been written in pencil on the label in an unknown hand.
Pierre s.n. is both better preserved than Thorel 1593 and shows the characters of the species more clearly. Therefore, it is the more appropriate choice as lectotype. Thorel 1593, which was collected in Ubon Ratchathani, might have been a more satisfactory choice, because it would explain the epithet “ubonense”, but it is a mixed collection (see under Eriocaulon nautiliforme above), which makes it less suitable for lectotypification. Thorel 1593 [P00224183] is a syntype of Eriocaulon ubonense.
Additional Notes on the Lectotype of Eriocaulon quinquangulare
Eriocaulon quinquangulare L., Sp. Pl. 87 (1753); Kunth, Enum. Pl. 3: 556 (1841); Hooker (1894: 582); Hooker in Trimen, Handb. Fl. Ceylon 5: 9 (1900); Ruhland in Engl., Pflanzenr. IV, 30 (Heft 13): 85 (Reference Ruhland and Engler1903); Lecomte, Fl. Indo-Chine 7(1): 12 (Reference Lecomte1912); Fyson, J. Indian Bot. 2(2–3): 31 (1923); P.H. Hộ, Ill. Fl. Vietnam 3(1): 396 (1993); C.D.K.Cook, Aquatic Wetland Pl. India: 199 (1996); S.M.Phillips in Dassan. & Clayton, Revis. Handb. Fl. Ceylon 11: 68 (Reference Phillips, Dassanayake and Clayton1997); Z.X.Zhang in U.Kück, Diss. Bot. 313: 111 (Reference Zhang and Kück1999); R.Ansari & N.P.Balakr., Fam. Eriocaul. India: 97–100 (Reference Ansari and Balakrishnan2009). – Type: CEYLON. P. Hermann s.n. (lectotype BM, Herb. Hermann 2: 21, no. 48, selected by Phillips, Reference Phillips, Dassanayake and Clayton1997).
Phillips (Reference Phillips, Dassanayake and Clayton1997) cited the type of this species as follows: “Type: Ceylon, Hermann s.n. (BM-SL)”. Our investigations have shown that three elements of original material survive, so a lectotype was required. Phillips's statement may be taken as a lectotypification, but because Hermann's material may be found scattered through some 15 volumes of the Sloane herbarium, and in a further five bound Hermann volumes not in the Sloane herbarium, we give a more precise reference here.
It is now possible to be certain that this species does not match any specimens collected in Cambodia, Laos and Vietnam. Eriocaulon quinquangulare is therefore a species of Sri Lanka and India that reaches western and north-western Thailand (Chiang Mai and Tak provinces) but no further east.
Acknowledgements
We should like to thank the curators of the herbaria at K and P for the loan of specimens while Amornrat Prajaksood was at TCD, and the curators of B, BR, K and P for the loan of specimens to Phetlasy Souladeth at E. This project was financially supported by a research studentship to Amornrat Prajaksood from Trinity College Dublin, September 2001–September 2004, and by a Darwin Fellowship to Phetlasy Souladeth, 2013–2014. We particularly thank Mark Spencer and Charlie Jarvis (BM) for their help in locating original material at BM, the Hermann collection and LINN; Sylvia Phillips (K) for her overview of the family; Mark Watson (E) for his knowledge of Buchanan-Hamilton; Hans-Joachim Esser (M) for his knowledge of Martius's collection; and David Middleton (SING) for helpful discussion of critical nomenclatural points.
