E D I N BU RG H J O U R N A L O F B O TA N Y
74 (1): 5–13 (2017)
5
© Trustees of the Royal Botanic Garden Edinburgh (2016)
doi: 10.1017/S0960428616000238
T Y P I F I C AT I O N O F NA M E S I N E R I O C AU LO N I N
T H E F LO R A O F T H A I L A N D A N D F LO R A O F
C A M B O D I A , L AO S A N D V I E T NA M
P. S o u la d e t h 1 , 2, A . P ra ja k s o o d 3 , 4, J. A . N. Pa r n e l l 4
& M . F. N e w m a n 2
Lectotypes are designated for eight names in Eriocaulon in tropical Asia, namely
E. alatum, E. hamiltonianum, E. hookerianum, E. infirmum, E. lanigerum, E. nautiliforme,
E. nigrum and E. ubonense. Additional information on the lectotype of Eriocaulon
quinquangulare is given.
Keywords. Eriocaulon, nomenclature, Southeast Asia, typification.
I n t ro d u c t i o n
Eriocaulon L. is a large genus of c.400 species distributed widely in the tropics and
subtropics (Stützel, 1998; Mabberley, 2008). 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 (1912) 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 (1908a,b), namely, Eriocaulon alatum Lecomte (1908b, p. 104), Eriocaulon lanigerum
Lecomte (1908b, p. 109), Eriocaulon nautiliforme Lecomte (1908a, p. 89), Eriocaulon
nigrum Lecomte (1908b, p. 107) and Eriocaulon ubonense Lecomte (1908b, p. 109).
1
2
3
4
Department of Forest Management, Faculty of Forestry Science, National University of Laos, PO Box
7322, Laos. E-mail: p.souladeth@nuol.edu.la
Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh EH3 5LR, Scotland, UK.
Applied Taxonomic Research Center, Department of Biology, Faculty of Science, Khon Kaen University,
Thailand.
Herbarium, Department of Botany and Trinity Centre for Biodiversity Research, School of Natural
Sciences, Trinity College Dublin, Dublin 2, Ireland
6
P. S O U L A D E T H E T A L .
Lecomte (1908a,b) 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., 2012). 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
(1832) and Eriocaulon infirmum Steud., Syn. Pl. Glumac. 2: 271 (1855), 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 (1829–1849) 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 (1829–1849) 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 (1829–1849) 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
T Y P I F I C AT I O N O F N A M E S I N E R I O C AU L O N
7
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., 2012).
L e c t o ty p i f i c at i o n
Eriocaulon alatum Lecomte, J. Bot. (Morot) sér. 2, 1: 104 (1908b) & Fl. Indo-Chine
7(1): 17 (1912); Merr., Fl. Manila: 136 (1912); 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, 1959),
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 (1999: 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 (1999: 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 (1832); Kunth,
Enum. Pl. 3: 552 (1841); Steud., Syn. Pl. Glumac. 2: 270 (1855); Körn., Linnaea
27: 679 (1856); Ruhland in Engl., Pflanzenr. IV, 30 (Heft 13): 107 (1903);
Z.X.Zhang in U.Kück, Diss. Bot. 313: 169–172 (1999); R.Ansari & N.P.Balakr.,
Fam. Eriocaul. India: 122 (2009). – Type: INDIA. Gualpara, F. Hamilton, s.n.
(lectotype BR [BR000000918773], designated here; isolectotypes, E [E00027135]; KW [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.
8
P. S O U L A D E T H E T A L .
r 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.
r 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.
r 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 (1894); Ruhland
in Engl., Pflanzenr. IV, 30 (Heft 13): 83 (1903); Lecomte Fl. Indo-Chine 7(1): 10
(1912); P.H. Hộ, Ill. Fl. Vietnam 3(1): 392 (1993); Beaman & R.S. Beaman (1998:
137); Z.X.Zhang in U.Kück, Diss. Bot. 313: 143 (1999); R.Ansari & N.P.Balakr.,
Fam. Eriocaul. India: 28 (2009). – 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 (1998) 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 (1998) use the modern place name spellings Kemburongoh and
T Y P I F I C AT I O N O F N A M E S I N E R I O C AU L O N
9
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 (1950), 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 (1998). 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 (2009) 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 (1855: 271); P.Royen, Nova Guinea 10(1–2): 38 (1959).
≡ Eriocaulon gracile Mart. in Wall., Pl. Asiat. Rar. 3: 29 (1832), non Eriocaulon gracile
Bong., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 1: 634 (1831).
– 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 (1855) published the name Eriocaulon infirmum for Eriocaulon gracile Martius
(1832). Although Steudel did not say so explicitly, his name Eriocaulon infirmum is
a replacement name, because Eriocaulon gracile Bongard (1831) had already been
published, based on a different type. Bongard’s species is from Brazil and is now called
Syngonanthus gracilis (Bong.) Ruhland.
Martius (1832: 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 (1908b); Fl. IndoChine 7(1): 8 (1912). – 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
10
P. S O U L A D E T H E T A L .
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., 2012).
Eriocaulon nautiliforme Lecomte, J. Bot. (Morot) sér. 2, 1: 89 (1908a); J. Bot. (Morot)
sér. 2, 1: 105–106 (1908b); Fl. Indo-Chine 7(1): 7 (1912). – Type: CAMBODIA.
Kampot, 6 December 1903, C. Geoffray 215 (lectotype P [P00224175], designated
here).
Lecomte (1908a) 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:
r Geoffray 215 [P00224175], collected at Kampot, Cambodia
r Pierre s.n. [P00224176], collected at Chaudoc, Vietnam
r Thorel 1593 [B 10 0086998], collected at Bassac, Laos [image seen at the Virtual
Herbarium Berolinense (Botanischer Garten und Botanisches Museum BerlinDahlem, no date)]
r Thorel 1593 [K, without barcode number], collected at Bassac, Laos
r 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 (1999) 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 highresolution 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).
T Y P I F I C AT I O N O F N A M E S I N E R I O C AU L O N
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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 (1908b); Fl. Indo-Chine 7(1):
16 (1912). – 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 (1908b); Fl. Indo-Chine
7(1): 11 (1912); 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.
r Pierre s.n. [P00224182]. The label states “[Cambodia 10/1874]” in a modern hand.
r 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.
A d d i t i o n a l N o t e s o n t h e L e c t o ty p e o f E R I O C AU L O N
Q U I N Q UA N G U L A R E
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 (1903); Lecomte, Fl. Indo-Chine 7(1): 12
(1912); 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 (1997); Z.X.Zhang in U.Kück, Diss.
Bot. 313: 111 (1999); R.Ansari & N.P.Balakr., Fam. Eriocaul. India: 97–100 (2009).
– Type: CEYLON. P. Hermann s.n. (lectotype BM, Herb. Hermann 2: 21, no. 48,
selected by Phillips, 1997).
12
P. S O U L A D E T H E T A L .
Phillips (1997) 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.
Ac k n ow l e d g e m e n t s
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.
References
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Received 14 September 2016; accepted for publication 22 September 2016;
first published online 17 November 2016