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Abstract

Following recent molecular phylogenetic studies in Old World Gesneriaceae the nomenclatural implications for names in Henckelia are examined. New combinations are made in Codonoboea and Loxocarpus to account for species now excluded from Henckelia. A list is presented in which the current position of all species hitherto assigned to Henckelia is given, including the new combination Henckelia rotundata (Barnett) D.J.Middleton & Mich.Möller. A new combination in Oreocharis is made.
EDINBURGH JOURNAL OF BOTANY 70 ( 3 ): 385 – 404 (2013)
© Trustees of the Royal Botanic Garden Edinburgh (2013)
doi:10.1017/S0960428613000127
385
THE CURRENT STATUS OF THE SPECIES
HITHERTO ASSIGNED TO HENCKELIA
(GESNERIACEAE)
D . J . M IDDLETON 1 , A . W EBER 2 , T . L . Y AO 3 , S . S ONTAG 2 &
M . M ÖLLER 1
Following recent molecular phylogenetic studies in Old World Gesneriaceae the
nomenclatural implications for names in Henckelia are examined. New combinations
are made in Codonoboea and Loxocarpus to account for species now excluded from
Henckelia . A list is presented in which the current position of all species hitherto
assigned to Henckelia is given, including the new combination Henckelia rotundata
(Barnett) D.J.Middleton & Mich.Möller. A new combination in Oreocharis is made.
Keywords . Codonoboea , Gesneriaceae , Henckelia , Loxocarpus , new combinations .
I NTRODUCTION
In recent years there has been a large number of changes in generic delimitation in
Gesneriaceae as a result of both detailed morphological study and insights from
molecular phylogenetic analyses (Möller et al. , 2009 , 2011a , 2011b ; Wei et al. , 2010 ;
Puglisi et al. , 2011 ; Wang et al. , 2011 ; Weber et al. , 2011a , 2011b , 2011c ). The study
by Weber et al. ( 2011a ) was ostensibly to clarify relationships in the heterogeneous
genus Chirita Buch.-Ham. ex D.Don but the conclusions also had significant ramifi-
cations in other genera, particularly in Henckelia Spreng. The objective of the current
paper is to clarify the nomenclatural consequences of changes in the delimitation of
Henckelia .
Henckelia was resurrected from synonymy of Didymocarpus Wall. by Weber & Burtt
(1998). It encompassed the species formerly placed in Didymocarpus sect. Orthoboea
(South India, Sri Lanka), the species of all (but one: Didymocarpus sect. Elati Ridl.)
sections of Didymocarpus from Peninsular Thailand and Malesia, as well as the
Malesian genera Codonoboea Ridl., Loxocarpus R.Br., and Platyadenia B.L.Burtt,
totalling about 180 species. The first molecular phylogenetic studies to include a range
of Henckelia species confirmed that these species indeed did not form a monophyletic
group with Didymocarpus but also began to suggest that the newly defined Henckelia
1 Royal Botanic Garden Edinburgh , 20A Inverleith Row , Edinburgh EH3 5LR , Scotland, UK . E-mail for
correspondence: d.middleton@rbge.ac.uk
2 University of Vienna , Faculty Centre of Biodiversity , Department of Structural and Functional Botany ,
Rennweg 14 , 1030 Vienna , Austria .
3 Forest Research Institute Malaysia , 52109 Kepong , Selangor , Malaysia .
D. J. MIDDLETON ET AL. 386
might also not be monophyletic (Möller et al. , 2009 ). Weber et al. ( 2011a ) showed
conclusively that the South Indian and Sri Lankan species of Henckelia formed a
monophyletic group with most species of Chirita sect. Chirita but that some species of
Chirita sect. Chirita and the other sections in the genus were only distantly related to
these. The type species of Henckelia , H. incana (Vahl) Spreng., is from South India.
Henckelia has priority over Chirita and the nomenclatural consequences for all spe-
cies previously assigned to Chirita were presented in Weber et al. ( 2011a ). The same
study also showed that all the Malesian species of Henckelia , by far the majority
of species in the genus, did not form a monophyletic group with the South Indian/
Sri Lankan species. The focus in Weber et al. ( 2011a ) was on the fate of species assigned
to Chirita but it was acknowledged there that the results and conclusions had huge
implications for our understanding of the genus Henckelia . Henckelia is still recog-
nised but with a delimitation considerably different from that used in a number of
other recent works (Weber & Burtt, 1998; Vitek et al. , 2000 ; Weber, 2004 ; Banka &
Kiew, 2009 ; Kiew, 2009 ). Before Weber et al. ( 2011a ) there were approximately 180 spe-
cies of Henckelia and afterwards about 56 species. However, only 14 species, namely
those of Henckelia sect. Henckelia (formerly Didymocarpus sect. Orthoboea Benth.)
from South India and Sri Lanka, are common to both delimitations.
Weber et al. ( 2011a ) noted that the correct generic name for most of the southern
Thai and Malesian species formerly placed in Henckelia is Codonoboea . The species of
Henckelia sections Didymanthus (C.B.Clarke) A.Weber & B.L.Burtt, Heteroboea
(Benth.) A.Weber & B.L.Burtt and Glossadenia A.Weber & B.L.Burtt (Weber &
Burtt, 1998) consistently form a well-supported monophyletic group and can confi-
dently all be assigned to Codonoboea . Kiew & Lim ( 2011 ) made combinations for all
the Peninsular Malaysian taxa and Kartonegoro ( 2012 ) has made a combination for
the single species known from Sulawesi. In addition Rafidah et al. ( 2011 ) have made
the combination Codonoboea elata (Ridl.) Rafidah for the misplaced Chirita elata
Ridl. Below we make combinations in Codonoboea for the remaining species in this
group from Peninsular Thailand and the rest of Malesia. We acknowledge that con-
siderably more basic taxonomic work is necessary in these areas to properly define the
species but we consider it essential that they have names in Codonoboea pending that
work so as to avoid conflicting nomenclatural frameworks.
A smaller number of species in Thailand and Malesia which were excluded from
Henckelia by Weber et al. ( 2011a ) did not group with those species now in Codonoboea .
These correspond to the species of Loxocarpus that had been synonymised into
Henckelia by Weber & Burtt (1998). The species of Loxocarpus are found in the Asian
twisted-fruited clade (using the terminology of Möller et al. , 2009 ) but doubts remain
as to whether the species form a monophyletic group (Yao, 2012 ). However, given
that Loxocarpus was explicitly excluded from the newly defined Henckelia by Weber
et al. ( 2011a ) the species must revert to Loxocarpus and await more research on their
relationships. The species have recently been revised by Yao ( 2012 ) and 20 named
species are presently recognised. Most of these 20 species already have combinations
in Loxocarpus but below we make new combinations where necessary.
CURRENT STATUS OF SPECIES HITHERTO ASSIGNED TO HENCKELIA 387
We make new combinations only for those taxa which, to the best of our knowl-
edge, are currently recognised. In the list below and in the list of all names ever pub-
lished in Henckelia in Appendix 1 we largely follow taxon delimitation in Weber &
Burtt (1998), Vitek et al. ( 2000 ), Weber et al. ( 2000 ), Burtt ( 2001 ), Kiew & Lim ( 2011 )
and Yao ( 2012 ).
We do not make combinations for the section names in Henckelia which were rec-
ognised by Weber & Burtt (1998) and which now belong to Codonoboea : sect.
Didymanthus , sect. Heteroboea , and sect. Glossadenia . We abstain from making the
formal transfers pending further research and simply list the appropriate sections
hitherto described in Didymocarpus and Henckelia in the synonymy of Codonoboea .
A new combination in Oreocharis is made in Appendix 2 .
N OMENCLATURAL CONSEQUENCES
Codonoboea
Codonoboea Ridl., Fl. Malay Penins. 2: 533 (1923). – Didymocarpus sect. Codonoboea
(Ridl.) Kiew, Blumea 35: 171 (1990). – Type: Codonoboea leucocodon (Ridl.) Ridl.,
designated by Ivanina ( 1967 ).
Didymocarpus sect. Heteroboea Benth. in Benth. & Hook.f., Gen. Pl. 2: 1022 (1876).
Henckelia sect. Heteroboea (Benth.) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen
70: 334 (1998 [‘1997’]). – Type: Didymocarpus crinitus (Jack) Spreng. ( Codonoboea
crinita (Jack) C.L.Lim).
Didymocarpus sect. Didymanthus C.B.Clarke in A.DC. & C.DC., Monogr. Phan.
5(1): 82 (1883). – Henckelia sect. Didymanthus (C.B.Clarke) A.Weber & B.L.Burtt,
Beitr. Biol. Pflanzen 70: 334 (1998 [‘1997’]). – Type: Didymocarpus serratus R.Br.
( Codonoboea serrata (R.Br.) A.Weber).
Didymocarpus sect. Kompsoboea C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1):
87 (1883). – Type: Didymocarpus kompsoboea C.B.Clarke ( Codonoboea komp-
soboea (C.B.Clarke) A.Weber).
Didymocarpus sect. Salicini Ridl., J. Linn. Soc. Bot. 32: 514 (1896). – Type:
Didymocarpus salicinus Ridl. ( Codonoboea salicina (Ridl.) C.L.Lim).
Didymocarpus sect. Reptantes Ridl., J. Straits Branch Roy. Asiat. Soc. 44: 29 (1905).
– Type: Didymocarpus reptans Jack ( Codonoboea reptans (Jack) C.L.Lim).
Paraboea sect. Campanulatae Ridl., J. Straits Branch Roy. Asiat. Soc. 44: 63 (1905).
– Type: Paraboea campanulata Ridl. (= Codonoboea hirta (Ridl.) Kiew).
Didymocarpus sect. Boeopsis Ridl., J. Straits Branch Roy. Asiat. Soc. 49: 22 (1907).
– Type: Didymocarpus heterophyllus Ridl. ( Codonoboea heterophylla (Ridl.)
C.L.Lim).
Didymocarpus sect. Pectinati Ridl., Fl. Malay Penins. 2: 508 (1923). – Type:
Didymocarpus pectinatus Oliv. ( Codonoboea pectinata (Oliv.) Kiew).
Platyadenia B.L.Burtt, Notes Roy. Bot. Gard. Edinburgh 31: 51 (1971). – Type:
Platyadenia descendens B.L.Burtt ( Codonoboea descendens (B.L.Burtt) A.Weber).
D. J. MIDDLETON ET AL. 388
Didymocarpus sect. Venusti Kiew, Malay. Nat. J. 48: 203 (1995) [‘ Venustus ’]. – Type:
Didymocarpus venustus Ridl. ( Codonoboea venusta (Ridl.) Kiew).
Henckelia sect. Glossadenia A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 335 (1998
[‘1997’]). – Type: Henckelia flavobrunnea (Ridl.) A.Weber ( Codonoboea flavo-
brunnea (Ridl.) Kiew).
Perennial herbs, caulescent or forming a rosette, occasionally creeping; usually some-
what woody. Leaves opposite or alternate, petiolate to sessile. Inflorescences axillary,
one- to few- to many-flowered, pedunculate, single-flowered or cymose, then flowers
in pairs; flowers 5-merous. Calyx of 5 lobes divided to the base. Corolla consisting of
a tube and 5 spreading lobes, 2-lipped with a 3-lobed lower lip, which is generally
longer, and a 2-lobed upper lip. Fertile stamens 2, anthers adnate face to face. Disc a
flat-topped or lobed ring, sometimes not a complete ring or lacking. Ovary cylindric,
style narrower, stigma peltate or clavate. Capsules plagiotropic, usually narrow and
long, dehiscing dorsally to form a gutter.
About 120 species from southern Thailand and throughout Malesia. The Peninsular
Malaysian species are enumerated in Kiew & Lim ( 2011 ) and the single Sulawesi spe-
cies by Kartonegoro ( 2012 ). Below are new combinations for the remaining species.
We have not indicated the type of type (holotype, lectotype, isotype, etc.) for all
names because for those that possibly lack a holotype, or have not yet been lectotyp-
ifed, this should be done as part of a detailed revision of the taxa.
Codonoboea alternifolia (C.B.Clarke) A.Weber, comb. nov. Didymocarpus alternifo-
lius C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 106 (1883). – Paraboea
alternifolia (C.B.Clarke) B.L.Burtt, Notes Roy. Bot. Gard. Edinburgh 24: 46
(1962). – Henckelia alternifolia (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70:
339 (1998 [‘1997’]). – Type: W Borneo, Kalimantan, Sebalouw, near Sambas, 15–19
x 1874, Teysmann 10779 (holo FI!). – Distribution: Borneo.
Codonoboea amoena (C.B.Clarke) A.Weber, comb. nov. Didymocarpus amoenus
C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 87 (1883). – Henckelia amoena
(C.B.Clarke) B.L.Burtt in Coode et al., Checkl. Fl. Pl. Gymnosperms Brunei
Darussalam 438 (1996). – Type: Borneo borealis, 1877–78, Burbidge s.n. (BM, K).
– Distribution: Borneo.
Codonoboea angustifolia (C.B.Clarke) A.Weber, comb. nov. Didymocarpus angustifolius
C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 89 (1883). – Henckelia angusti-
folia (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 339 (1998 [‘1997’]). – Type:
Borneo, Sarawak, in monte Mattang, Beccari 3629 (FI-B, K). – Distribution: Borneo.
Codonoboea appressipilosa (B.L.Burtt) D.J.Middleton, comb. nov. Henckelia appres-
sipilosa B.L.Burtt, Thai Forest Bull., Bot. 29: 98 (2001). – Type: Thailand,
Narathiwat, Sungai Kolok, Nikhom Waeng, 2 iii 1974, K. Larsen & S. Larsen
32803 (holo AAU). – Distribution: Thailand.
CURRENT STATUS OF SPECIES HITHERTO ASSIGNED TO HENCKELIA 389
Codonoboea bakoensis (B.L.Burtt) A.Weber, comb. nov. Didymocarpus bakoensis
B.L.Burtt, Notes Roy. Bot. Gard. Edinburgh 36: 152 (1978). – Henckelia bakoensis
(B.L.Burtt) B.L.Burtt, Beitr. Biol. Pflanzen 70: 340 (1998 [‘1997’]). – Type: Sarawak,
First Division, Bako National Park, Telok Asam, 17 v 1962, Burtt & Woods 1839
(holo E; iso SAR). – Distribution: Borneo.
Codonoboea bakoensis (B.L.Burtt) A.Weber var. tenuior (C.B.Clarke) A.Weber,
comb. nov. Didymocarpus leptocalyx C.B.Clarke var. tenuior C.B.Clarke in A.DC. &
C.DC., Monogr. Phan. 5(1): 91 (1883). – Henckelia bakoensis (B.L.Burtt) B.L.Burtt
var. tenuior (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 340 (1998 [‘1997’]).
– Type: Borneo, prope Kuteing [Kuching], Beccari (holo FI-B). – Distribution:
Borneo.
Codonoboea battamensis (Ridl.) A.Weber, comb. nov. Didymocarpus battamensis
Ridl., J. Straits Branch Roy. Asiat. Soc. 49: 22 (1908). – Henckelia battamensis
(Ridl.) B.L.Burtt, Beitr. Biol. Pflanzen 70: 340 (1998 [‘1997’]). – Type: Pulau
Battam, an island south of Singapore, ix 1905, Kloss s.n. (SING). – Distribution:
Batam Island (Indonesia).
Codonoboea beccarii (C.B.Clarke) A.Weber, comb. nov. Didymocarpus beccarii
C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 89 (1883). – Henckelia beccarii
(C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 340 (1998 [‘1997’]). – Syntypes:
Sarawak, in colle Sakarang, Beccari 3884 (FI-B); in colle Cumpang, v 1867, Beccari
3442 (FI-B, K); Kapuas, Teysmann in herb. Beccari 8441 (FI-B). – Distribution:
Borneo.
Codonoboea bullata (C.B.Clarke) A.Weber, comb. nov. Didymocarpus bullatus
C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 92 (1883). – Henckelia bullata
(C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 339 (1998 [‘1997’]). – Syntypes:
Borneo, Sarawak, 750 m, Lobb s.n. (K); Beccari 2108 (FI-B). – Distribution:
Borneo.
Codonoboea corniculata (Jack) D.J.Middleton, comb. nov. Didymocarpus cornicula-
tus Jack, Malayan Misc. I, 2: 4 (1820). – Henckelia corniculata (Jack) Spreng.,
Syst. Veg., ed. 16, 4(2): 13 (1827). – Type: Sumatra, Tappanooly, Jack s.n. (G).
– Distribution: Sumatra.
Codonoboea corrugata (Mendum) D.J.Middleton, comb. nov. Henckelia corrugata
Mendum, Edinburgh J. Bot. 58: 439 (2001). – Type: Philippines, Palawan, side of
ravine above Camp 3 towards Cleopatra’s Needle, 940 m, 22 i 1998, Mendum et al.
25558 (holo PNH; iso E). – Distribution: Philippines (Palawan).
Codonoboea crenata (Baker) A.Weber, comb. nov. Didymocarpus crenatus Baker,
Kew Bull. 1896: 25 (1896). – Henckelia crenata (Baker) B.L.Burtt in Coode et al.,
Checkl. Fl. Pl. Gymnosperms Brunei Darussalam 438 (1996). – Type: Borneo, Sabah,
Sandakan, iv 1895, Creagh (holo K). – Distribution: Borneo.
D. J. MIDDLETON ET AL. 390
Codonoboea dentata (Ridl.) A.Weber, comb. nov. Didymocarpus dentatus Ridl.,
J. Linn. Soc. 32: 515 (1896). – Henckelia dentata (Ridl.) B.L.Burtt, Beitr. Biol.
Pflanzen 70: 343 (1998 [‘1997’]). – Type: Indonesia, Lingga, 1893, Hullett s.n.
(SING). – Distribution: Lingga Islands (Indonesia).
Codonoboea descendens (B.L.Burtt) A.Weber, comb. nov. Platyadenia descendens
B.L.Burtt, Notes Roy. Bot. Gard. Edinburgh 31: 51 (1971). – Henckelia descendens
(B.L.Burtt) B.L.Burtt, Beitr. Biol. Pflanzen 70: 343 (1998 [‘1997’]). – Type: Sarawak,
First Division, Bako National Park, Telok Delima, 19 v 1962, Burtt & Woods
B.1856 (holo E; iso SAR). – Distribution: Borneo.
Codonoboea elegans (C.B.Clarke) A.Weber, comb. nov. Didymocarpus elegans
C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 88 (1883). – Henckelia elegans
(C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 344 (1998 [‘1997’]). – Type:
Borneo, 1853, Lobb s.n. (holo K). – Distribution: Borneo.
Codonoboea filicalyx (B.L.Burtt) D.J.Middleton, comb. nov. Henckelia filicalyx
B.L.Burtt, Thai Forest Bull., Bot. 29: 95 (2001). – Type: Thailand, Narathiwat,
Nikhom Waeng, 4 iii 1974, K. Larsen & S. Larsen 32914 (holo AAU; iso E).
– Distribution: Thailand.
Codonoboea follicularis (C.B.Clarke) A.Weber, comb. nov. Didymocarpus follicularis
C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 98 (1883). – Henckelia follicu-
laris (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 345 (1998 [‘1997’]). – Type:
Borneo, Karimata, Teysmann in herb. Beccari 11218 (FI-B; photo E). – Distribution:
Borneo.
Codonoboea gracilipes (C.B.Clarke) A.Weber, comb. nov. Didymocarpus gracilipes
C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 97 (1883). – Henckelia gra-
cilipes (C.B.Clarke) B.L.Burtt in Coode et al., Checkl. Fl. Pl. Gymnosperms Brunei
Darussalam 438 (1996). – Type: Borneo, Sarawak, Beccari 3229 (FI-B, G, K, M, P, W).
– Distribution: Borneo.
Codonoboea koerperi (B.L.Burtt) A.Weber, comb. nov. Didymocarpus koerperi
B.L.Burtt, Notes Roy. Bot. Gard. Edinburgh 47: 236 (1990). – Henckelia koerperi
(B.L.Burtt) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 347 (1998 [‘1997’]).
– Type: Cultivated plant originally from Sumatra, Pajakumbuh, Harau valley,
c.800 m, Körper s.n. (holo E; iso M, WU). – Distribution: Sumatra.
Codonoboea kolokensis (B.L.Burtt) D.J.Middleton, comb. nov. Henckelia kolokensis
B.L.Burtt, Thai Forest Bull., Bot. 29: 95 (2001). – Type: Thailand, Narathiwat,
Sungai Kolok, Nikhom Waeng, 4 iii 1974, K. Larsen & S. Larsen 32913 (holo AAU;
iso E). – Distribution: Thailand.
Codonoboea kompsoboea (C.B.Clarke) A.Weber, comb. nov. Didymocarpus komp-
soboea C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 92, tab. 10 (1883).
Henckelia kompsoboea (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 347
CURRENT STATUS OF SPECIES HITHERTO ASSIGNED TO HENCKELIA 391
(1998 [‘1997’]). – Type: Borneo, ‘Sarawak, in colle Kumpang’, Beccari 3440 (holo
FI-B). – Distribution: Borneo.
Codonoboea lanceolata (C.B.Clarke) A.Weber, comb. nov. Didymocarpus lanceola-
tus C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 89 (1883). – Henckelia
lanceolata (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 348 (1998 [‘1997’]).
– Type: ‘Borneo boreali-orientalis; in montibus’, Lobb s.n. (holo K). – Distribution:
Borneo.
Codonoboea leptocalyx (C.B.Clarke) A.Weber, comb. nov. Didymocarpus lepto-
calyx C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 91 (1883). – Henckelia
leptocalyx (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 348 (1998 [‘1997’]).
– Syntypes: Borneo, Sarawak, 750 m, Beccari 1794 (FI-B, G-DC, K, P); Lobb s.n.
(K). – Distribution: Borneo.
Codonoboea murutorum (B.L.Burtt) A.Weber, comb. nov. Didymocarpus muruto-
rum B.L.Burtt, Notes Roy. Bot. Gard. Edinburgh 36: 153 (1978). – Henckelia
murutorum (B.L.Burtt) B.L.Burtt, Beitr. Biol. Pflanzen 70: 350 (1998 [‘1997’]).
– Type: Sarawak, Fifth Division, route from Bakelalan to Gunung Murud above
Sungei Konap, c.1320 m, 23 ix 1967, Burtt & Martin B.5172 (holo E; iso SAR).
– Distribution: Borneo.
Codonoboea myricifolia (Ridl.) A.Weber, comb. nov. Didymocarpus myricifolius
Ridl., J. Straits Branch Roy. Asiat. Soc. 44: 53 (1905). – Henckelia myricifolia
(Ridl.) B.L.Burtt in Coode et al., Checkl. Fl. Pl. Gymnosperms Brunei Darussalam
438 (1996). – Type: Borneo, Sabah, Bongaya, Labuk Bay, Ridley s.n. (SING).
– Distribution: Borneo.
Codonoboea nervosa (C.B.Clarke) A.Weber, comb. nov. Didymocarpus nervosus
C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 95 (1883). – Henckelia nervosa
(C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 350 (1998 [‘1997’]). – Type:
Borneo, Sarawak, in colle Sakarang, Beccari 3889 (holo FI-B). – Distribution:
Borneo.
Codonoboea pagonensis (B.L.Burtt) A.Weber, comb. nov. Henckelia pagonensis
B.L.Burtt in Coode et al., Checkl. Fl. Pl. Gymnosperms Brunei Darussalam 437
(1996). – Type: Brunei, Temburong, Bukit Pagon, east ridge, 1470 m, 19 vii 1990,
K.M. Wong 1784 (holo BRUN). – Distribution: Borneo.
Codonoboea pleuropogon (B.L.Burtt) A.Weber, comb. nov. Didymocarpus pleuropo-
gon B.L.Burtt, Notes Roy. Bot. Gard. Edinburgh 31: 44 (1971). – Henckelia pleuro-
pogon (B.L.Burtt) B.L.Burtt in Coode et al., Checkl. Fl. Pl. Gymnosperms Brunei
Darussalam 438 (1996). – Type: Cultivated plant grown on from Burtt & Martin
B.5050, originally from Sarawak, Third Division, SE Hose Mts., hill E of overhang
at Bukit Semako, 16 viii 1967, Royal Botanic Garden Edinburgh cultivated specimen
C.5639 (holo E). – Distribution: Borneo.
D. J. MIDDLETON ET AL. 392
Codonoboea porphyrea (B.L.Burtt) D.J.Middleton, comb. nov. Henckelia porphyrea
B.L.Burtt, Thai Forest Bull., Bot. 29: 96 (2001). – Type: Thailand, Ranong,
Khao Pota Luang Kaeo, 22 vi 1974, Geesink et al. 7417 (holo BKF; iso AAU).
– Distribution: Thailand.
Codonoboea punctata (C.B.Clarke) A.Weber, comb. nov. Didymocarpus punctatus
C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 90 (1883). – Henckelia punc-
tata (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 353 (1998 [‘1997’]). – Type:
‘Borneo boreali-orientalis, in ripis fluminum’, Lobb s.n. (holo K). – Distribution:
Borneo.
Codonoboea racemosa (Jack) A.Weber, comb. nov. Didymocarpus racemosus Jack,
Trans. Linn. Soc. 14: 34 (1823). – Henckelia racemosa (Jack) Spreng., Syst. Veg.,
ed. 16, 4(2): 13 (1827). – Type: Sumatra, west coast, at Tappanooly, Jack s.n. (prob-
ably lost). – Distribution: Sumatra.
Codonoboea reticulosa (C.B.Clarke) A.Weber, comb. nov. Didymocarpus reticulosus
C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 90 (1883). – Henckelia reticu-
losa (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 354 (1998 [‘1997’]). – Syntypes:
Borneo, Sarawak, 750 m, 1865–68, Beccari 1713 (FI-B); 1865–68, Beccari 2067 (FI-B,
G-DC, K, P); Lobb (K). – Distribution: Borneo.
Codonoboea scabrinervia (C.B.Clarke) A.Weber, comb. nov. Didymocarpus scabri-
nervius C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 91 (1883). – Henckelia
scabrinervia (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 355 (1998 [‘1997’]).
– Syntypes: Borneo, ‘Sarawak, in monte Poe, alt. 1400 m’, Beccari 2423 (FI-B);
Montrado, Teysmann 10883 (FI-B, L). – Distribution: Borneo.
Codonoboea serrata (R.Br.) A.Weber, comb. nov. Didymocarpus serratus R.Br.,
Cyrtandreae 119 (1839). – Henckelia serrata (R.Br.) A.Weber & B.L.Burtt, Beitr.
Biol. Pflanzen 70: 356 (1998 [‘1997’]). – Type: Sumatra?, Horsfield s.n. (BM).
– Distribution: probably Sumatra.
Codonoboea simplex (Kraenzl.) A.Weber, comb. nov. Didymocarpus simplex Kraenzl.,
Mitt. Inst. Allg. Bot. Hamburg 7: 88 (1927). – Henckelia simplex (Kraenzl.)
B.L.Burtt, Beitr. Biol. Pflanzen 70: 356 (1998 [‘1997’]). – Type: West-Borneo
[Kalimantan], ‘zwischen dem Bukit Raja und dem Bukit Menabai, um 1000 m’, 25
xii 1924, Winkler 1058 (HBG; photo E). – Distribution: Borneo.
Codonoboea teres (C.B.Clarke) A.Weber, comb. nov. Didymocarpus teres C.B.Clarke
in A.DC. & C.DC., Monogr. Phan. 5(1): 88 (1883). – Henckelia teres (C.B.Clarke)
B.L.Burtt, Beitr. Biol. Pflanzen 70: 357 (1998 [‘1997’]). – Syntypes: Borneo,
‘Sarawak, prope Kuteing’ [Kuching], Beccari 385 (FI-B, K); Beccari 1353 (FI-B).
– Distribution: Borneo.
Codonoboea virginea (B.L.Burtt) A.Weber, comb. nov. Didymocarpus virgineus
B.L.Burtt, Notes Roy. Bot. Gard. Edinburgh 36: 154 (1978). – Henckelia virginea
CURRENT STATUS OF SPECIES HITHERTO ASSIGNED TO HENCKELIA 393
(B.L.Burtt) B.L.Burtt, Beitr. Biol. Pflanzen 70: 358 (1998 [‘1997’]). – Type: Borneo,
Sarawak, Fifth Division, route from Bakelelan to Gunung Murud, ridge above
Sungai Bor, c.4°N, 115°38 E, c.1320 m, 25 ix 1967, Burtt & Martin B.5203 (holo E;
iso SAR). – Distribution: Borneo.
Codonoboea vulcanica (Ridl.) A.Weber, comb. nov. Didymocarpus vulcanicus
Ridl., J. Malayan Branch Roy. Asiat. Soc. 1: 80 (1923). – Henckelia vulcanica
(Ridl.) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 359 (1998 [‘1997’]). – Type:
Sumatra, Bandar Bharu, Gunong Sibayak, 6 viii 1921, Nur 7314 (K). – Distribution:
Sumatra.
Codonoboea woodii (Merr.) A.Weber, comb. nov. Didymocarpus woodii Merr.,
Philipp. J. Sci. 26: 489 (1925). – Henckelia woodii (Merr.) A.Weber & B.L.Burtt,
Beitr. Biol. Pflanzen 70: 359 (1998 [‘1997’]). – Type: Philippines, Palawan,
Malampaya Bay, 8 x 1922, Merrill 11587 (G, K, L, P, PNH, US, W). – Distribution:
Philippines.
Loxocarpus
Loxocarpus R.Br., Cyrtandreae 120 (1839). – Didymocarpus Wall. sect. Loxocarpus
(R.Br.) Benth. in Benth. & Hook.f., Gen. Pl. 2: 1022 (1876). – Roettlera Vahl sect.
Loxocarpus (R.Br.) Fritsch in Nat. Pflanzenfam. 4, 36: 147 (1894). – Henckelia
Spreng. sect. Loxocarpus (R.Br.) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70:
334 (1998). – Type: Loxocarpus incanus R.Br.
Rosette plants with or without rootstock, sometimes creeping, rarely caulescent.
Indumentum usually of a layer of silky hairs, generally whitish, sometimes buff, usu-
ally dense, rarely sparse on lamina upper surface. Leaves petiolate. Inflorescence a
reduced cyme with a single flower, or cymose, pedunculate. Flowers 5-merous. Calyx
lobes free almost to base. Corolla usually campanulate or flat-faced, rarely shortly
tubular; lobes more or less isomorphic but lower three lobes usually distinguishably
larger than upper two lobes, upper lobes rarely fused. Disc usually absent, sometimes
vestigial. Stamens 2; anthers coherent, usually included within corolla tube in cam-
panulate and shortly tubular flowers or exposed in flat-faced flowers. Ovary conical,
usually short and thick at base, sometimes slender; stigma punctiform, sometimes
capitate and rarely peltate. Capsules plagiocarpic, often ovoid or conical, sometimes
slender and gutter shaped, usually straight or slightly curved, rarely corniculate,
dehiscing dorsally to form a splash cup or gutter.
More than 20 species in southern Thailand and western Malesia.
Loxocarpus coodei (B.L.Burtt) T.L.Yao, comb. nov. Henckelia coodei B.L.Burtt in
Coode et al., Checkl. Fl. Pl. Gymnosperms Brunei Darussalam 437 (1996). – Type:
Brunei, Temburong, Temburong River at Wong Nguan rapids, 120 m, 5 iii 1990,
Coode 6617 (holo K; iso BRUN).
D. J. MIDDLETON ET AL. 394
Loxocarpus incanus R.Br. var. sekayuensis (Banka & Kiew) T.L.Yao, comb. et stat.
nov. Henckelia sekayuensis Banka & Kiew, Edinburgh J. Bot. 66: 254 (2009).
– Type: Peninsular Malaysia, Terengganu, Sekayu Recreational Forest, 29 viii 1986,
Anthonysamy SA 638 (holo KEP).
Loxocarpus sericiflavus (Kiew & Banka) T.L.Yao, comb. nov. Henckelia sericiflava
Kiew & Banka, Edinburgh J. Bot. 66: 18 (2009). – Type: Peninsular Malaysia,
Johor, Gunung Panti, 5 xii 1936, Corner SFN 32538 (holo SING; iso SAR).
Loxocarpus taeniophyllus (B.L.Burtt) T.L.Yao, comb. nov. Henckelia taeniophylla
B.L.Burtt in Coode et al., Checkl. Fl. Pl. Gymnosperms Brunei Darussalam 438
(1996). – Type: Brunei, Temburong, Bangar, northern slope of Bkt. Bangar, 10–
100 m, 18 i 1964, Hotta 13263 (holo KYO; iso E).
Loxocarpus violoides (C.B.Clarke) T.L.Yao, comb. nov. Didymocarpus violoides
C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 97 (1883). – Roettlera violoides
(C.B.Clarke) Kuntze, Rev. Gen. Pl. 2: 477 (1891). – Henckelia violoides (C.B.Clarke)
B.L.Burtt in Coode et al., Checkl. Fl. Pl. Gymnosperms Brunei Darussalam 438
(1996). – Type: NE of Borneo, Lobong Peak, 5000 ft, Lobb s.n. (holo K).
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Received 19 June 2012 ; accepted for publication 10 December 2012
A PPENDIX 1
Present placement of species referred to Henckelia . Current name in bold . For further
nomenclatural details, where necessary, see Weber & Burtt (1998), Weber et al. ( 2000 ),
Vitek et al. ( 2000 ), Burtt ( 2001 ), Kiew & Lim ( 2011 ) and Yao (2012 ).
Henckelia adenocalyx (Chatterjee) D.J.Middleton & Mich.Möller, Taxon 60(3): 774 (2011).
Henckelia alata (Roxb.) A.Dietr., Sp. Pl., ed. 6, 1: 573 (1831). – Torenia alata (Roxb.) Benth.
Henckelia alba (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 338 (1998 [‘1997’]). – Codonoboea alba
(Ridl.) C.L.Lim
Henckelia albomarginata (Hemsl.) A.Weber, Beitr. Biol. Pflanzen 70: 338 (1998 [‘1997’]).
Codonoboea albomarginata (Hemsl.) Kiew
Henckelia alternans (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 338 (1998 [‘1997’]). – Codonoboea
alternans (Ridl.) Kiew
Henckelia alternifolia (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 339 (1998 [‘1997’]).
Codonoboea alternifolia (C.B.Clarke) A.Weber
D. J. MIDDLETON ET AL. 396
Henckelia amoena (C.B.Clarke) B.L.Burtt in Coode et al., Checkl. Fl. Pl. Gymnosperms Brunei
Darussalam 438 (1996). – Codonoboea amoena (C.B.Clarke) A.Weber
Henckelia anachoreta (Hance) D.J.Middleton & Mich.Möller, Taxon 60(3): 774 (2011).
Henckelia angusta (C.B.Clarke) D.J.Middleton & Mich.Möller, Taxon 60(3): 774 (2011).
Henckelia angustifolia (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 339 (1998 [‘1997’]).
Codonoboea angustifolia (C.B.Clarke) A.Weber
Henckelia anthonyi (Kiew) A.Weber, Beitr. Biol. Pflanzen 70: 339 (1998 [‘1997’]). – Codonoboea
anthonyi (Kiew) C.L.Lim
Henckelia anthonysamyi Banka, Edinburgh J. Bot. 66(2): 249 (2009). – Synonym of Loxocarpus
sericiflavus (Kiew & Banka) T.L.Yao
Henckelia appressipilosa B.L.Burtt, Thai Forest Bull., Bot. 29: 98 (2001). – Codonoboea
appressipilosa (B.L.Burtt) D.J.Middleton
Henckelia argentea (B.L.Burtt) B.L.Burtt, Beitr. Biol. Pflanzen 70: 339 (1998 [‘1997’]). – Loxocarpus
argenteus B.L.Burtt
Henckelia aromatica (D.Don) Spreng., Syst. Veg., ed. 16, 4(2): 13 (1827). – Didymocarpus aro-
maticus D.Don
Henckelia ascendens (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 339 (1998 [‘1997’]). – Codonoboea
ascendens (Ridl.) C.L.Lim
Henckelia atrosanguinea (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 340 (1998 [‘1997’]).
Codonoboea atrosanguinea (Ridl.) C.L.Lim
Henckelia auriculata (J.M.Li & S.X.Zhu) D.J.Middleton & Mich.Möller, Taxon 60(3): 774 (2011).
Henckelia bakoensis (B.L.Burtt) B.L.Burtt, Beitr. Biol. Pflanzen 70: 340 (1998 [‘1997’]).
Codonoboea bakoensis (B.L.Burtt) A.Weber
Henckelia bakoensis var. tenuior (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 340 (1998
[‘1997’]). – Codonoboea bakoensis (B.L.Burtt) A.Weber var. tenuior (C.B.Clarke) A.Weber
Henckelia barbata (Jack) Spreng., Syst. Veg., ed. 16, 4(2): 13 (1827). = Liebigia speciosa (Blume)
Endl.
Henckelia battamensis (Ridl.) B.L.Burtt, Beitr. Biol. Pflanzen 70: 340 (1998 [‘1997’]).
Codonoboea battamensis (Ridl.) A.Weber
Henckelia beccarii (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 340 (1998 [‘1997’]).
Codonoboea beccarii (C.B.Clarke) A.Weber
Henckelia bifolia
(D.Don) A.Dietr., Sp. Pl., ed. 6, 1: 574 (1831).
Henckelia bombycina (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 340 (1998 [‘1997’]). – Codonoboea
bombycina (Ridl.) C.L.Lim
Henckelia breviflora (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 340 (1998 [‘1997’]). – Codonoboea
breviflora (Ridl.) Kiew
Henckelia briggsioides (W.T.Wang) D.J.Middleton & Mich.Möller, Taxon 60(3): 775 (2011).
Henckelia browniana A.Weber, Beitr. Biol. Pflanzen 70: 341 (1998 [‘1997’]). – Loxocarpus
incanus R.Br.
Henckelia bullata (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 341 (1998 [‘1997’]).
Codonoboea bullata (C.B.Clarke) A.Weber
Henckelia burttii D.J.Middleton & Mich.Möller, Taxon 60(3): 775 (2011).
Henckelia caelestis (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 341 (1998 [‘1997’]). – Codonoboea
caelestis Ridl.
Henckelia caerulea (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 341 (1998 [‘1997’]). – Loxocarpus
caeruleus (Ridl.) Ridl.
Henckelia calcarea (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 341 (1998 [‘1997’]). – Codonoboea
calcarea (Ridl.) Kiew
Henckelia calva (C.B.Clarke) D.J.Middleton & Mich.Möller, Taxon 60(3): 775 (2011).
Henckelia capensis A.Braun ex C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 151 (1883),
nom. inval.
CURRENT STATUS OF SPECIES HITHERTO ASSIGNED TO HENCKELIA 397
Henckelia castaneifolia (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 341 (1998 [‘1997’]). – Codonoboea
castaneifolia (Ridl.) Kiew
Henckelia caulescens (B.L.Burtt) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 342 (1998
[‘1997’]). – Loxocarpus caulescens B.L.Burtt
Henckelia ceratoscyphus (B.L.Burtt) D.J.Middleton & Mich.Möller, Taxon 60(3): 775 (2011).
Henckelia cinerea (D.Don) Spreng., Syst. Veg., ed. 16, 4(2): 14 (1827). – Didymocarpus cinereus
D.Don
Henckelia codonion (Kiew) A.Weber, Beitr. Biol. Pflanzen 70: 342 (1998 [‘1997’]). – Codonoboea
codonion (Kiew) C.L.Lim
Henckelia communis (Gardner) D.J.Middleton & Mich.Möller, Taxon 60(3): 775 (2011).
Henckelia conicapsularis (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 342 (1998 [‘1997’]).
Loxocarpus conicapsularis (C.B.Clarke) B.L.Burtt
Henckelia coodei B.L.Burtt in Coode et al., Checkl. Fl. Pl. Gymnosperms Brunei Darussalam
437 (1996). – Loxocarpus coodei (B.L.Burtt) T.L.Yao
Henckelia corneri (Kiew) A.Weber, Beitr. Biol. Pflanzen 70: 342 (1998 [‘1997’]). – Codonoboea
corneri (Kiew) Kiew
Henckelia corniculata (Jack) Spreng., Syst. Veg., ed. 16, 4(2): 18 (1827). – Codonoboea corniculata
(Jack) D.J.Middleton
Henckelia corrugata Mendum, Edinburgh J. Bot. 58(3): 439 (2001). – Codonoboea corrugata
(Mendum) D.J.Middleton
Henckelia craspedodroma (Kiew) A.Weber, Beitr. Biol. Pflanzen 70: 342 (1998 [‘1997’]).
Codonoboea craspedodroma (Kiew) Kiew
Henckelia crenata (Baker) B.L.Burtt in Coode et al., Checkl. Fl. Pl. Gymnosperms Brunei
Darussalam 438 (1996). – Codonoboea crenata (Baker) A.Weber
Henckelia crinita (Jack) Spreng., Syst. Veg., ed. 16, 4(2): 18 (1827). – Codonoboea crinita (Jack)
C.L.Lim
Henckelia crocea (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 343 (1998 [‘1997’]). – Codonoboea
crocea (Ridl.) C.L.Lim
Henckelia curtisii (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 343 (1998 [‘1997’]). – Codonoboea
curtisii (Ridl.) C.L.Lim
Henckelia davisonii (Kiew) A.Weber, Beitr. Biol. Pflanzen 70: 343 (1998 [‘1997’]). – Codonoboea
davisonii (Kiew) Kiew
Henckelia dawnii (Kiew) A.Weber, Beitr. Biol. Pflanzen 70: 343 (1998 [‘1997’]). –
Codonoboea
dawnii (Kiew) Kiew
Henckelia densifolia (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 343 (1998 [‘1997’]). – Codonoboea
densifolia (Ridl.) C.L.Lim
Henckelia dentata (Ridl.) B.L.Burtt, Beitr. Biol. Pflanzen 70: 343 (1998 [‘1997’]). – Codonoboea
dentata (Ridl.) A.Weber
Henckelia descendens (B.L.Burtt) B.L.Burtt, Beitr. Biol. Pflanzen 70: 343 (1998 [‘1997’]).
Codonoboea descendens (B.L.Burtt) A.Weber
Henckelia dibangensis (B.L.Burtt, S.K.Srivast. & Mehrotra) D.J.Middleton & Mich.Möller,
Taxon 60(3): 775 (2011).
Henckelia dielsii (Borza) D.J.Middleton & Mich.Möller, Taxon 60(5): 1525 (2011).
Henckelia diffusa B.L.Burtt in Coode et al., Checkl. Fl. Pl. Gymnosperms Brunei Darussalam
437 (1996). – Synonym of Loxocarpus violoides (C.B.Clarke) T.L.Yao
Henckelia dimidiata (Wall. ex C.B.Clarke) D.J.Middleton & Mich.Möller, Taxon 60(3): 775
(2011).
Henckelia doryphylla (B.L.Burtt) A.Weber, Beitr. Biol. Pflanzen 70: 344 (1998 [‘1997’]).
Codonoboea doryphylla (B.L.Burtt) C.L.Lim
Henckelia elegans (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 344 (1998 [‘1997’]).
Codonoboea elegans (C.B.Clarke) A.Weber
D. J. MIDDLETON ET AL. 398
Henckelia elongata (Jack) Spreng., Syst. Veg., ed. 16, 4(2): 18 (1827). – Didissandra elongata
(Jack) C.B.Clarke
Henckelia ericiflora (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 344 (1998 [‘1997’]). – Codonoboea
ericiflora (Ridl.) Ridl.
Henckelia ericii A.Weber, Beitr. Biol. Pflanzen 70: 344 (1998 [‘1997’]). – Loxocarpus holttumii
M.R.Hend.
Henckelia falcata (Kiew) A.Weber, Beitr. Biol. Pflanzen 70: 344 (1998 [‘1997’]). – Codonoboea
falcata (Kiew) Kiew
Henckelia fasciata (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 344 (1998 [‘1997’]). – Codonoboea
fasciata (Ridl.) C.L.Lim
Henckelia fasciculiflora (W.T.Wang) D.J.Middleton & Mich.Möller, Taxon 60(3): 775 (2011).
Henckelia filicalyx B.L.Burtt, Thai Forest Bull., Bot. 29: 95 (2001). – Codonoboea filicalyx
(B.L.Burtt) D.J.Middleton
Henckelia fischeri (Gamble) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 344 (1998
[‘1997’]).
Henckelia flava (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 344 (1998 [‘1997’]). – Codonoboea flava
(Ridl.) Kiew
Henckelia flavescens (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 344 (1998 [‘1997’]). – Codonoboea
flavescens (Ridl.) Kiew
Henckelia flavobrunnea (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 344 (1998 [‘1997’]). – Codonoboea
flavobrunnea (Ridl.) Kiew
Henckelia floccosa (Thwaites) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 344 (1998
[‘1997’]).
Henckelia floribunda (M.R.Hend.) A.Weber, Beitr. Biol. Pflanzen 70: 345 (1998 [‘1997’]).
Codonoboea floribunda (M.R.Hend.) C.L.Lim
Henckelia follicularis (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 345 (1998 [‘1997’]).
Codonoboea follicularis (C.B.Clarke) A.Weber
Henckelia forrestii (J.Anthony) D.J.Middleton & Mich.Möller, Taxon 60(3): 775 (2011).
Henckelia frutescens (Jack) Spreng., Syst. Veg., ed. 16, 4(2): 18 (1827). – Didissandra frutescens
(Jack) C.B.Clarke
Henckelia fruticola (H.W.Li) D.J.Middleton & Mich.Möller, Taxon 60(3): 775 (2011).
Henckelia gambleana (C.E.C.Fisch.) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 345 (1998
[‘1997’]).
Henckelia gardneri B.L.Burtt in Coode et al., Checkl. Fl. Pl. Gymnosperms Brunei Darussalam
437 (1996). – Synonym of Loxocarpus repens B.L.Burtt
Henckelia geitleri (A.Weber) A.Weber, Beitr. Biol. Pflanzen 70: 345 (1998 [‘1997’]). – Codonoboea
geitleri (A.Weber) C.L.Lim
Henckelia glabrata (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 345 (1998 [‘1997’]). – Codonoboea
glabrata (Ridl.) Kiew
Henckelia gracilipes (C.B.Clarke) B.L.Burtt in Coode et al., Checkl. Fl. Pl. Gymnosperms
Brunei Darussalam 438 (1996). – Codonoboea gracilipes (C.B.Clarke) A.Weber
Henckelia grandiflora (Retz.) A.Dietr., Sp. Pl., ed. 6, 1: 573 (1831). – Synonym of Lindernia
antipoda (L.) Alston
Henckelia grandifolia A.Dietr., Sp. Pl., ed. 6, 1: 576 (1831).
Henckelia heterophylla (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 346 (1998 [‘1997’]). – Codonoboea
heterophylla (Ridl.) C.L.Lim
Henckelia heterostigma (B.L.Burtt) D.J.Middleton & Mich.Möller, Taxon 60(3): 775 (2011).
Henckelia hirsuta (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 346 (1998 [‘1997’]). – Codonoboea
hirsuta (Ridl.) C.L.Lim
Henckelia hirta (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 346 (1998 [‘1997’]). – Codonoboea hirta
(Ridl.) Kiew
CURRENT STATUS OF SPECIES HITHERTO ASSIGNED TO HENCKELIA 399
Henckelia hispida (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 346 (1998 [‘1997’]). – Codonoboea
hispida (Ridl.) Kiew
Henckelia holttumii (M.R.Hend.) A.Weber, Beitr. Biol. Pflanzen 70: 346 (1998 [‘1997’]).
Codonoboea holttumii (M.R.Hend.) C.L.Lim
Henckelia hookeri (C.B.Clarke) D.J.Middleton & Mich.Möller, Taxon 60(3): 775 (2011).
Henckelia humboldtiana (Gardner) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 346 (1998
[‘1997’]).
Henckelia humilis (Miq.) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 347 (1998 [‘1997’]).
Codonoboea humilis (Miq.) A.Weber
Henckelia inaequalis (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 347 (1998 [‘1997’]). – Codonoboea
inaequalis (Ridl.) Kiew
Henckelia incana (Vahl) Spreng., Syst. Veg., ed. 16, 1: 38 (1824).
Henckelia infundibuliformis (W.T.Wang) D.J.Middleton & Mich.Möller, Taxon 60(3): 775
(2011).
Henckelia innominata (B.L.Burtt) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 347 (1998
[‘1997’]).
Henckelia insignis (C.B.Clarke) D.J.Middleton & Mich.Möller, Taxon 60(3): 776 (2011).
Henckelia johorica (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 347 (1998 [‘1997’]). – Codonoboea
johorica (Ridl.) Kiew
Henckelia kelantanensis Kiew, Gard. Bull. Singapore 61(1): 73 (2009). – Codonoboea kelantan-
ensis (Kiew) Kiew
Henckelia kjellbergii B.L.Burtt, Beitr. Biol. Pflanzen 70: 378 (1998 [‘1997’]). – Codonoboea kjell-
bergii (B.L.Burtt) Karton. [Kartonegoro, 2012 ].
Henckelia koerperi (B.L.Burtt) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 347 (1998
[‘1997’]). – Codonoboea koerperi (B.L.Burtt) A.Weber
Henckelia kolokensis B.L.Burtt, Thai Forest Bull., Bot. 29: 95 (2001). – Codonoboea kolokensis
(B.L.Burtt) D.J.Middleton
Henckelia kompsoboea (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 347 (1998 [‘1997’]).
Codonoboea kompsoboea (C.B.Clarke) A.Weber
Henckelia lacei (W.W.Sm.) D.J.Middleton & Mich.Möller, Taxon 60(3): 776 (2011).
Henckelia lachenensis (C.B.Clarke) D.J.Middleton & Mich.Möller, Taxon 60(3): 776 (2011).
Henckelia lanceolata (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 348 (1998 [‘1997’]).
Codonoboea lanceolata (C.B.Clarke) A.Weber
Henckelia lancifolia (M.R.Hend.) A.Weber, Beitr. Biol. Pflanzen 70: 348 (1998 [‘1997’]).
Codonoboea lancifolia (M.R.Hend.) C.L.Lim
Henckelia leiophylla (Kiew) A.Weber, Beitr. Biol. Pflanzen 70: 348 (1998 [‘1997’]). –
Codonoboea
leiophylla (Kiew) C.L.Lim
Henckelia leptocalyx (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 348 (1998 [‘1997’]).
Codonoboea leptocalyx (C.B.Clarke) A.Weber
Henckelia leucantha (Kiew) A.Weber, Beitr. Biol. Pflanzen 70: 348 (1998 [‘1997’]). – Codonoboea
leucantha (Kiew) Kiew
Henckelia leucocodon (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 348 (1998 [‘1997’]). – Codonoboea
leucocodon (Ridl.) Ridl.
Henckelia lilacina (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 348 (1998 [‘1997’]). – Codonoboea
lilacina (Ridl.) Ridl.
Henckelia longipedicellata (B.L.Burtt) D.J.Middleton & Mich.Möller, Taxon 60(3): 776
(2011).
Henckelia longipes (C.B.Clarke) A.Weber, Beitr. Biol. Pflanzen 70: 349 (1998 [‘1997’]). – Codonoboea
longipes (C.B.Clarke) Kiew
Henckelia longipetiolata (B.L.Burtt) B.L.Burtt, Beitr. Biol. Pflanzen 70: 349 (1998 [‘1997’]).
– Synonym of Loxocarpus stapfii (Kraenzl.) B.L.Burtt
D. J. MIDDLETON ET AL. 400
Henckelia longisepala (H.W.Li) D.J.Middleton & Mich.Möller, Taxon 60(3): 776 (2011).
Henckelia lyrata (Wight) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 349 (1998 [‘1997’]).
Henckelia macrophylla (D.Don) Spreng., Syst. Veg., ed. 16, 4(2): 14 (1827). – Didymocarpus
macrophyllus D.Don
Henckelia macrostachya (E.Barnes) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 349 (1998
[‘1997’]).
Henckelia malayana (Hook.f.) A.Weber, Beitr. Biol. Pflanzen 70: 349 (1998 [‘1997’]). – Codonoboea
malayana (Hook.f.) Kiew
Henckelia marginata (C.B.Clarke) A.Weber, Beitr. Biol. Pflanzen 70: 349 (1998 [‘1997’]).
Codonoboea marginata (C.B.Clarke) C.L.Lim
Henckelia meeboldii (W.W.Sm. & Ramaswami) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70:
349 (1998 [‘1997’]).
Henckelia meijeri (B.L.Burtt) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 350 (1998 [‘1997’]).
Loxocarpus meijeri B.L.Burtt
Henckelia miniata (Kiew) A.Weber, Beitr. Biol. Pflanzen 70: 350 (1998 [‘1997’]). – Codonoboea
miniata (Kiew) C.L.Lim
Henckelia minima (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 350 (1998 [‘1997’]). – Synonym
of Loxocarpus incanus R.Br.
Henckelia mishmiensis (Debb. ex Biswas) D.J.Middleton & Mich.Möller, Taxon 60(3): 776
(2011).
Henckelia missionis (Wall. ex R.Br.) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 350 (1998
[‘1997’]).
Henckelia modesta (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 350 (1998 [‘1997’]). – Codonoboea
modesta (Ridl.) Kiew
Henckelia monantha (W.T.Wang) D.J.Middleton & Mich.Möller, Taxon 60(3): 776 (2011).
Henckelia monophylla (C.B.Clarke) D.J.Middleton & Mich.Möller, Taxon 60(3): 776 (2011).
Henckelia moonii (Gardner) D.J.Middleton & Mich.Möller, Taxon 60(3): 776 (2011).
Henckelia murutorum (B.L.Burtt) B.L.Burtt, Beitr. Biol. Pflanzen 70: 350 (1998 [‘1997’]).
Codonoboea murutorum (B.L.Burtt) A.Weber
Henckelia myricifolia (Ridl.) B.L.Burtt in Coode et al., Checkl. Fl. Pl. Gymnosperms Brunei
Darussalam 438 (1996). – Codonoboea myricifolia (Ridl.) A.Weber
Henckelia nana A.Weber, Beitr. Biol. Pflanzen 70: 350 (1998 [‘1997’]). – Codonoboea pumila
(Ridl.) C.L.Lim
Henckelia nervosa (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 350 (1998 [‘1997’]).
Codonoboea nervosa (C.B.Clarke) A.Weber
Henckelia nitida (Kiew & A.Weber) A.Weber, Beitr. Biol. Pflanzen 70: 351 (1998 [‘1997’]).
Codonoboea nitida (Kiew & A.Weber) Kiew
Henckelia nivea (Kiew) A.Weber, Beitr. Biol. Pflanzen 70: 351 (1998 [‘1997’]). – Codonoboea
nivea Kiew
Henckelia oblonga (D.Don) Spreng., Syst. Veg., ed. 16, 4(2): 13 (1827). – Didymocarpus
oblongus D.Don
Henckelia oblongifolia (Roxb.) D.J.Middleton & Mich.Möller, Taxon 60(3): 776 (2011).
Henckelia oppositifolia (Retz.) A.Dietr., Sp. Pl., ed. 6, 1: 572 (1831). – Synonym of Lindernia
hyssopioides (L.) Haines
Henckelia ovalifolia (Wight) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 351 (1998
[‘1997’]).
Henckelia pagonensis B.L.Burtt in Coode et al., Checkl. Fl. Pl. Gymnosperms Brunei
Darussalam 437 (1996). – Codonoboea pagonensis (B.L.Burtt) A.Weber
Henckelia papillosa (M.R.Hend.) A.Weber, Beitr. Biol. Pflanzen 70: 351 (1998 [‘1997’]). Notes:
Banka & Kiew ( 2009 ) and Yao ( 2012 ) excluded Henckelia papillosa from Loxocarpus and
said its affinities are with the species now placed in Codonoboea. However, Kiew & Lim
CURRENT STATUS OF SPECIES HITHERTO ASSIGNED TO HENCKELIA 401
( 2011 ) did not refer to the species in their Codonoboea paper. The status of the name, there-
fore, remains uncertain.
Henckelia parviflora (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 352 (1998 [‘1997’]). – Codonoboea
parviflora (Ridl.) Kiew
Henckelia pauziana Kiew, Gard. Bull. Singapore 61(1): 74 (2009). – Codonoboea pauziana
(Kiew) Kiew
Henckelia pectinata (Oliv.) A.Weber, Beitr. Biol. Pflanzen 70: 352 (1998 [‘1997’]). – Codonoboea
pectinata (Oliv.) Kiew
Henckelia peduncularis (B.L.Burtt) D.J.Middleton & Mich.Möller, Taxon 60(3): 776 (2011).
Henckelia petiolaris (C.B.Clarke) B.L.Burtt in Coode et al., Checkl. Fl. Pl. Gymnosperms
Brunei Darussalam 438 (1996). – Synonym of Loxocarpus verbeniflos (C.B.Clarke)
B.L.Burtt
Henckelia platypus (C.B.Clarke) A.Weber, Beitr. Biol. Pflanzen 70: 352 (1998 [‘1997’]).
Codonoboea platypus (C.B.Clarke) C.L.Lim
Henckelia pleuropogon (B.L.Burtt) B.L.Burtt in Coode et al., Checkl. Fl. Pl. Gymnosperms
Brunei Darussalam 438 (1996). – Codonoboea pleuropogon (B.L.Burtt) A.Weber
Henckelia plicata (D.Don) Spreng., Syst. Veg., ed. 16, 4(2): 14 (1827). – Synonym of
Didymocarpus macrophyllus D.Don
Henckelia polyanthoides (Kiew) A.Weber, Beitr. Biol. Pflanzen 70: 352 (1998 [‘1997’]).
Codonoboea polyanthoides (Kiew) C.L.Lim
Henckelia porphyrea B.L.Burtt, Thai Forest Bull., Bot. 29: 96 (2001). – Codonoboea porphyrea
(B.L.Burtt) D.J.Middleton
Henckelia primulacea (C.B.Clarke) D.J.Middleton & Mich.Möller, Taxon 60(3): 776 (2011).
Henckelia primulifolia (D.Don) Spreng., Syst. Veg., ed. 16, 4(2): 14 (1827). – Didymocarpus primu-
lifolius D.Don
Henckelia primulina (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 352 (1998 [‘1997’]). – Codonoboea
primulina (Ridl.) Kiew
Henckelia procumbens B.L.Burtt, Beitr. Biol. Pflanzen 70: 353 (1998 [‘1997’]). – Loxocarpus
repens B.L.Burtt
Henckelia puerensis (Y.Y.Qian) D.J.Middleton & Mich.Möller, Taxon 60(3): 776 (2011).
Henckelia pulchella (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 353 (1998 [‘1997’]). – Codonoboea
pulchella (Ridl.) C.L.Lim
Henckelia pumila (D.Don) A.Dietr., Sp. Pl., ed. 6, 1: 574 (1831).
Henckelia punctata (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 353 (1998 [‘1997’]).
Codonoboea punctata (C.B.Clarke) A.Weber
Henckelia puncticulata (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 353 (1998 [‘1997’]).
Codonoboea puncticulata (Ridl.) C.L.Lim
Henckelia pycnantha (W.T.Wang) D.J.Middleton & Mich.Möller, Taxon 60(3): 776 (2011).
Henckelia pyroliflora (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 353 (1998 [‘1997’]). – Codonoboea
pyroliflora (Ridl.) Kiew
Henckelia quinquevulnera (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 353 (1998 [‘1997’]).
Codonoboea quinquevulnera (Ridl.) C.L.Lim
Henckelia racemosa (Jack) Spreng., Syst. Veg., ed. 16, 4(2): 13 (1827). – Codonoboea racemosa
(Jack) A.Weber
Henckelia ramosa (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 353 (1998 [‘1997’]). – Codonoboea
ramosa (Ridl.) Kiew
Henckelia repens (Bedd.) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 354 (1998 [‘1997’]).
Henckelia reptans (Jack) Spreng., Syst. Veg., ed. 16, 4(2): 14 (1827). – Codonoboea reptans
(Jack) C.L.Lim
Henckelia reptans var. monticola (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 354 (1998 [‘1997’]).
Codonoboea reptans var. monticola (Ridl.) C.L.Lim
D. J. MIDDLETON ET AL. 402
Henckelia reticulosa (C.B.Clarke) A.Weber, Beitr. Biol. Pflanzen 70: 354 (1998 [‘1997’]).
Codonoboea reticulosa (C.B.Clarke) A.Weber
Henckelia rexii (Hook.) Heynh., Nomencl. Bot. Hort. 1: 390 (1840). – Streptocarpus rexii
(Hook.) Lindl.
Henckelia ridleyana A.Weber, Beitr. Biol. Pflanzen 70: 354 (1998 [‘1997’]). – Codonoboea
ridleyana (B.L.Burtt) Kiew
Henckelia robinsonii (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 354 (1998 [‘1997’]). – Codonoboea
robinsonii (Ridl.) Kiew
Henckelia rotundata (Barnett) D.J.Middleton & Mich.Möller, comb. nov. Chirita rotun-
data Barnett, Nat. Hist. Bull. Siam Soc. 20: 16 (1961). – Type: Thailand, Chiang Mai,
Me Chem, 11 vii 1922, Kerr 6274 (lecto K, designated by Barnett ( 1961b ); iso ABD,
BKF, BM). This combination was included in Weber et al. ( 2011a ) but the basionym was
cited as having been published in Barnett ( 1961b ) rather than the slightly earlier Barnett
( 1961a ).
Henckelia roxburghiana A.Dietr., Sp. Pl., ed. 6, 1: 572 (1831). – Synonym of Lindernia ruel-
lioides (Colsm.) Pennell
Henckelia rubiginosa (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 355 (1998 [‘1997’]). – Codonoboea
rubiginosa (Ridl.) C.L.Lim
Henckelia rufescens (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 355 (1998 [‘1997’]).
Loxocarpus rufescens (C.B.Clarke) B.L.Burtt
Henckelia rugosa (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 355 (1998 [‘1997’]). – Codonoboea
rugosa (Ridl.) C.L.Lim
Henckelia salicina (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 355 (1998 [‘1997’]). – Codonoboea
salicina (Ridl.) C.L.Lim
Henckelia salicinoides (Kiew) A.Weber, Beitr. Biol. Pflanzen 70: 355 (1998 [‘1997’]).
Codonoboea salicinoides (Kiew) C.L.Lim
Henckelia scabrinervia (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 355 (1998 [‘1997’]).
Codonoboea scabrinervia (C.B.Clarke) A.Weber
Henckelia scortechinii (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 356 (1998 [‘1997’]). – The
current status of this name is unknown (see Kiew & Lim ( 2011 ) for further discussion)
Henckelia sekayuensis Banka & Kiew, Edinburgh J. Bot. 66(2): 254 (2009). – Loxocarpus incanus
var. sekayuensis (Banka & Kiew) T.L.Yao
Henckelia semitorta (C.B.Clarke) A.Weber, Beitr. Biol. Pflanzen 70: 356 (1998 [‘1997’]).
Loxocarpus semitortus (C.B.Clarke) Ridl.
Henckelia sericea (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 356 (1998 [‘1997’]). – Loxocarpus
sericeus (Ridl.) B.L.Burtt
Henckelia sericiflava Kiew & Banka, Edinburgh J. Bot. 66(2): 256 (2009). – Loxocarpus serici-
flavus (Kiew & Banka) T.L.Yao
Henckelia serrata (R.Br.) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 356 (1998 [‘1997’]).
Codonoboea serrata (R.Br.) A.Weber
Henckelia serratifolia (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 356 (1998 [‘1997’]). – Codonoboea
serratifolia (Ridl.) Kiew
Henckelia shuii (Z.Yu Li) D.J.Middleton & Mich.Möller, Taxon 60(3): 776 (2011).
Henckelia simplex (Kraenzl.) B.L.Burtt, Beitr. Biol. Pflanzen 70: 356 (1998 [‘1997’]). – Codonoboea
simplex (Kraenzl.) A.Weber
Henckelia smitinandii (B.L.Burtt) D.J.Middleton & Mich.Möller, Taxon 60(3): 776 (2011).
Henckelia soldanella (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 356 (1998 [‘1997’]). – Codonoboea
soldanella (Ridl.) C.L.Lim
Henckelia speciosa (Kurz) D.J.Middleton & Mich.Möller, Taxon 60(3): 777 (2011).
Henckelia stapfii (Kraenzl.) B.L.Burtt, Beitr. Biol. Pflanzen 70: 356 (1998 [‘1997’]). – Loxocarpus
stapfii (Kraenzl.) B.L.Burtt
CURRENT STATUS OF SPECIES HITHERTO ASSIGNED TO HENCKELIA 403
Henckelia stenophylla A.Weber, Beitr. Biol. Pflanzen 70: 357 (1998 [‘1997’]). – Loxocarpus
angustifolius Ridl.
Henckelia stolonifera (Kiew) A.Weber, Beitr. Biol. Pflanzen 70: 357 (1998 [‘1997’]). – Codonoboea
stolonifera (Kiew) Kiew
Henckelia taeniophylla B.L.Burtt in Coode et al., Checkl. Fl. Pl. Gymnosperms Brunei
Darussalam 438 (1996). – Loxocarpus taeniophyllus (B.L.Burtt) T.L.Yao
Henckelia tahanica (B.L.Burtt) A.Weber, Beitr. Biol. Pflanzen 70: 357 (1998 [‘1997’]). – Codonoboea
grandifolia (Ridl.) Kiew
Henckelia teres (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 357 (1998 [‘1997’]). – Codonoboea
teres (C.B.Clarke) A.Weber
Henckelia tibetica (Franch.) D.J.Middleton & Mich.Möller, Taxon 60(3): 777 (2011).
Henckelia tiumanica (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 357 (1998 [‘1997’]). – Codonoboea
tiumanica (Ridl.) C.L.Lim
Henckelia tunkui (Kiew) A.Weber, Beitr. Biol. Pflanzen 70: 357 (1998 [‘1997’]). – Loxocarpus
tunkui Kiew
Henckelia urticifolia (D.Don) A.Dietr., Sp. Pl., ed. 6, 1: 574 (1831).
Henckelia urticoides A.Weber, Beitr. Biol. Pflanzen 70: 358 (1998 [‘1997’]). – Codonoboea urti-
coides (A.Weber) Kiew
Henckelia venusta (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 358 (1998 [‘1997’]). – Codonoboea
venusta (Ridl.) Kiew
Henckelia verbeniflos (C.B.Clarke) B.L.Burtt, Beitr. Biol. Pflanzen 70: 358 (1998 [‘1997’]).
Loxocarpus verbeniflos (C.B.Clarke) B.L.Burtt
Henckelia villosa (D.Don) Spreng., Syst. Veg., ed. 16, 4(2): 13 (1827). – Didymocarpus villosus
D.Don
Henckelia violoides (C.B.Clarke) B.L.Burtt in Coode et al., Checkl. Fl. Pl. Gymnosperms Brunei
Darussalam 438 (1996). – Loxocarpus violoides (C.B.Clarke) T.L.Yao
Henckelia virginea (B.L.Burtt) B.L.Burtt, Beitr. Biol. Pflanzen 70: 358 (1998 [‘1997’]).
Codonoboea virginea (B.L.Burtt) A.Weber
Henckelia viscida (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70: 359 (1998 [‘1997’]). – Codonoboea
viscida (Ridl.) Kiew
Henckelia vulcanica (Ridl.) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 359 (1998 [‘1997’]).
Codonoboea vulcanica (Ridl.) A.Weber
Henckelia walkerae (Gardner) D.J.Middleton & Mich.Möller, Taxon 60(3): 777 (2011).
Henckelia wallichiana A.Dietr., Sp. Pl., ed. 6, 1: 574 (1831). – Synonym of Henckelia urticifolia
(D.Don.) A.Dietr.
Henckelia wightii (C.B.Clarke) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 359 (1998
[‘1997’]).
Henckelia woodii (Merr.) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 359 (1998 [‘1997’]).
Codonoboea woodii (Merr.) A.Weber
Henckelia yongii (Kiew) A.Weber, Beitr. Biol. Pflanzen 70: 359 (1998 [‘1997’]). – Codonoboea
yongii (Kiew) C.L.Lim
Henckelia zeylanica (R.Br.) A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 359 (1998 [‘1997’]).
D. J. MIDDLETON ET AL. 404
A PPENDIX 2
A new combination in Oreocharis
Many new combinations in Oreocharis were published by Möller et al. ( 2011b ). It has
come to our attention that the combination Oreocharis muscicola in that paper was
not validly made. We validate it here.
Oreocharis muscicola (Diels) Mich.Möller & A.Weber, comb. nov. Didissandra mus-
cicola Diels, Notes Roy. Bot. Gard. Edinburgh 5: 225 (1912). – Briggsia muscicola
(Diels) Craib, Notes Roy. Bot. Gard. Edinburgh 11: 264 (1919). – Type: China,
Yunnan, Mekong-Salwin [Salween] divide, vii 1905, Forrest 5095 (holo E).
Distribution: SW China, Bhutan, NE India, Burma.
... Henckelia belongs to the family Gesneriaceae and consists of about 76 species distributed in Sri Lanka, S and NE India, Nepal, Bhutan, S and SW China, N Vietnam, N Laos, and N Thailand (Weber et al. 2013, Middleton et al. 2013, Möller et al. 2017, Sirimongkol et al. 2019, Borah et al. 2019, Janeesha & Nampy 2020, Taram et al. 2021, Shi & Yang 2021. With 38 taxa, Henckelia is the largest gesneriaceous genus in India (Möller et al. 2017, Krishna & Lakshminarasimhan 2018, Borah et al. 2019, Janeesha & Nampy 2020. ...
... Recent molecular and morphology-based analyses demonstrated that the traditionally-defined Oreocharis was phylogenetically intertwined with nine previously defined small genera and acaulescent, rosette forming members of Briggsia Craib. The re-circumscribed Oreocharis is a strongly supported monophyletic group and placed in the subfamily Didymocarpoideae (Möller et al. 2011;Middleton et al. 2013). Since its re-definition in 2011, Oreocharis s.l. ...
Article
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Oreocharisphuongii, a new species of Gesneriaceae from central Vietnam, is described and illustrated here. The new species is most similar to O.longifolia by sharing peduncles up to 22 cm long, bracts 2, zygomorphic, yellow flowers with tubular corolla, stamens 4 with two pairs of coherent anthers and capsules up to 6 cm long. It mainly differs from the latter by the combination of some morphological characters of leaves (shape, base, apex and margin), size of calyx lobes, indumentum of corolla tube and inner surface of three lower corolla lobes. Detailed morphological description together with colour illustration, information on phenology, distribution, ecology, preliminarily conservation status of the new species and comparison with its similar species are also presented.
... The genus Henckelia Spreng., currently includes 75 species (Taram et al., 2021;Weber et al., 2011) distributed through Himalayas, Northeast and South India, Sri Lanka, Thailand, Laos and Vietnam (Middleton et al., 2013;Möller et al., 2017). India is represented by 39 species of which 22 occurs in Northeast India (Taram et al., 2021, revised). ...
Article
Herein, we present the first record of Henckelia longipedicellata (B.L.Burtt) D.J.Middleton & Mich.Möller (Gesneriaceae) for the flora of India based on specimens from Mayodia, Mishmi Hills, Lower Dibang Valley district of Arunachal Pradesh, India and its rediscovery after type collection. This species is easily recognized due to its solitary flowers with longer peduncles, deeply divided calyx lobes and ventrally pouched campanulate corolla. H. longipedicellata is so far known from Myanmar. A brief description along with additional notes and photographs are provided.
... Chirita Wood (1974: 131) [excluding species belonging to Damrongia Kerr ex Craib (1918: 364)] and the monotypic genus Hemiboeopsis W.T. Wang (1984: 397). After intensive field investigations in recent years, the re-circumscribed Henckelia currently comprises 74 species, which are distributed in Sri Lanka, India, Bangladesh, Nepal, Bhutan, S China, Myanmar, N Vietnam, N Laos, and N Thailand (Weber et al. 2011, Middleton et al. 2013, Janeesha & Nampy 2020, Nampy et al.2021, Shi & Yang 2021. In China, a total of 25 species of this genus have been recorded, with four species recently described, i.e., H. xinpingensis Y.H. Tan & Bin Yang in Yang et al. (2019: 196), H. nanxiheensis Lei Cai & Z.L. Dao in Cai et al. (2019: 152), H. multinervia Lei Cai & Z.L. Dao in Cai et al. (2019: 155) and H. connata X.Z. ...
Article
Full-text available
Henckelia inaequalifolia, a new species in the genus Henckelia of the family Gesneriaceae is described here. The detailed morphological description, photographic illustration and geographical distribution of the new species are provided. Its morphological affinities between related species are also discussed in this paper.
... has recently received great interests from taxonomists, with no less than 17 new species described after its recircumscription (Weber et al. 2011). At present, the genus comprises ca 70 species, most of which occurre in the Himalayas and Hengduan mountains while some others extend to Southeast Asia, Sri Lanka and south India (Weber et al. 2011, Middleton et al. 2013, Janeesha and Nampy 2020. Southwest China harbors a high diversity of Henckelia, and a total of 25 species have been recorded here. ...
Article
Full-text available
According to morphological observation and literature consultation, a new species of Henckelia (Gesneriaceae), H. connata X. Z. Shi & Li H. Yang, is described and illustrated. This new species resembles H. dimidiata, but differs in its dilated and connate petiole bases (versus separated), longer peduncles (4–10 cm long versus 0.3–2.2 cm long) and pedicels (1.5–4.0 cm long versus up to 1 cm long), lanceolate to triangular bracts (versus ovate), of smaller size (4–7 × 3–5 mm versus up to 16 × 13 mm), bluish purple to light blue corolla tube (versus yellow) and pubescence on most organs (glabrous or sparsely pubescent).
... Nearly 100 species mostly from southern India, Sri Lanka, southern Thailand and Malesia were transferred to three reinstated genera, viz. Henckelia Sprengel (1817: 402), Codonoboea Ridley (1923: 533) and Loxocarpus Brown (1838: 120) (Weber & Burtt 1998;Middleton et al. 2013), while a few other species were transferred to Streptocarpus Lindley (1828Lindley ( : t. 1173) (two Madagascan species) (Weber & Burtt 1998;Nishii et al. 2015), Petrocodon Hance (1883: 167) (three species from China and one species from northeastern Vietnam and northern Thailand) Li et al. 2016), and Tribounia D.J. Middleton in Middleton & Möller (2012: 1287 (one species from Thailand) respectively. With more than 20 species described mostly from Thailand and China since 1998, the newly circumscribed Didymocarpus currently consists of about 100 species distributed from India, Bangladesh, Nepal, Bhutan, southern China, Myanmar, Thailand, Vietnam, Cambodia, Laos, Peninsular Malaysia to Sumatra (Nangngam & Maxwell 2013;Nangngam & Middleton 2014;Möller 2019;Souvannakhoummane et al. 2019;Yang et al. 2019;POWO 2020;Adhikari & Möller 2020;Xie et al. 2020;Tran et al. 2020). ...
Article
Full-text available
Didymocarpus longicalyx, a new species of Gesneriaceae from southwestern Yunnan Province, China is described and illustrated based on detailed morphological comparisons with all known species of the genus Didymocarpus from both China and neighbouring countries. Morphologically, it is most similar to D. medogensis and D. glandulosus, but can be distinguished by its puberulent peduncles, calyx with ca. 8 mm long segments and claret corollas with three yellow spots and a few yellowish striations on upper lip.
... At least 30 new taxa of Oreocharis Benth. (Gesneriaceae) have been described and officially published (e.g., Cai et al. 2017Cai et al. , 2019Do et al. 2017;Chen et al. 2018;Guo et al. 2018;Möller et al. 2018;Pan et al. 2019;Yang et al. 2019) after the generic redefinition based on new evidence following the development of molecular phylogenetics in 2011 (Möller et al. 2011) and several later adjustments of the species (Middleton et al. 2013;Chen et al. 2014;Möller et al. 2014;Möller 2015). Oreocharis s.l. ...
Article
Full-text available
Two new species of Gesneriaceae, Oreocharis aimodisca and O. longipedicellata, from the limestone area of Yunnan Province, China, are described and illustrated. Their morphological relationship with similar species is discussed and colour photographs, detailed descriptions, distribution and habitat, as well as the IUCN endangered status are provided. We also discuss the accuracy of the scientific names of the described species O. tetrapterus from Guangxi, China in 2019 and O. brachypodus from Guizhou, China, in 2015, and put forward corrections related to name form.
Article
A new species of Henckelia (Gesneriaceae) is here described and illustrated. It is morphologically most similar to H. oblongifolia but differs in having lanceolate calyx lobes, two prominent longitudinal flaps on the inner surface of the corolla tube, bilobed stigma and tomentose capsules. A detailed description of the new species with colour photographs, affinities and a comparison with related species is given. Based on the present data, the new species is provisionally assessed as Critically Endangered (CR) according to IUCN Red List Categories and Criteria.
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The large, tropical island of Borneo has some of the world’s richest habitats for plant life, but faces increasing pressures from anthropogenic activities that threaten its biodiversity. With a good portion of the Bornean flora not critically studied, a comprehensive documentation of the numerous endemic taxa expected for the island is not yet complete. It is not known what the relative significance of endemic genera is compared to Bornean centres of endemism documented or predicted through modelling, and if they can inform current conservation plans. As a first step, we here present a synopsis of the endemic genera of Borneo, based on a comprehensive study of literature, herbarium specimens and distributional data, and an investigation of whether the genera have been included in molecular phylogenetic studies that confirm their monophyly. Such a review is timely since many generic delimitations have been shaped by molecular evidence used to test morphology-based taxonomy, while botanical collection and revisionary efforts continue. Our findings suggest that 65 vascular plant genera from 25 families may be considered endemic to Borneo. More than two-thirds (48) of these genera have had at least one species included in molecular phylogenetic studies, but of these, only 39 have been sufficiently sampled to be considered monophyletic with high confidence, or they are monotypic. Slightly over half (38) of the endemic genera are herbaceous. A majority of the genera have fruits or seeds specialised for dispersal by abiotic vectors, or unspecialised seeds. Almost two-thirds (42) of the endemic genera are monotypic, and some of these could represent relict lineages. We expect the current list of endemic genera to be relatively stable and aligned with recent taxonomic concepts, and that it serves to illuminate an interesting aspect of Borneo’s unique assemblage of endemic species.
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Based on molecular studies, the small Chinese genus Petrocodon (two species and one variety) has been recently enlarged to include the monotypic genera Calcareoboea, Paralagarosolen and Tengia. It is shown here that the (6-7) species of Lagarosolen, the monotypic Dolicholoma, a few species of Didymocarpus, and a number of new species that have recently been published (but not formally described) under Petrocodon and Lagarosolen should be included in this genus. This raises the size of the genus from five to around 20 species. With respect to the floral diversity (corolla form, size, and coloration; with the exception of Tengia, the androecium is always diandrous) and inferred pollination syndromes (different forms of melittophily, ornithophily, psycho- and/or sphingophily), Petrocodon represents one of the most varied genera of Old World Gesneriaceae, comparable to some New World genera.
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Five species are reinstated in Codonoboea Ridl. and new combinations are made for 74 species that occur in Peninsular Malaysia that were formerly included in Henckelia Spreng. Codonoboea albina (Ridl.) Kiew is reinstated at specific rank and var. winkleri (Ridl.) Kiew as a variety of C. malayana (Ridl.) Kiew. Lectotypes are designated where appropriate.
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Based on molecular data and a morphological evaluation, evidence is provided that the species of eleven, mostly small-sized and monotypic genera of Chinese Gesneriaceae (Ancylostemon, Bournea, Briggsia s.str., Dayaoshania, Deinocheilos, Isometrum, Opithandra, Oreocharis, Paraisometrum, Thamnocharis, Tremacron) form a highly-supported group in which the species interrelationships run across traditional generic boundaries. The data confirm previous doubts on the naturalness of some of these genera and, after a detailed discussion of the particular genera, the conclusion is reached that the whole group is best regarded as a single genus, Oreocharis, which is thus expanded to comprise over 80 species. A list of the species is given and the necessary transfers are made. The new delimitation provides a framework for studying the species relationships and working out an infrageneric classification. Oreocharis provides an excellent example of a major monophyletic group that has experienced a rapid radiation early in its evolution and shows manifold convergences in floral characters (corolla form and coloration, fertility of stamens, anther shape and dehiscence mode), apparently reflecting different pollination strategies, but has little variation in vegetative habit and fruit structure.
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A new species of Rhynchoglossum, R. capsulare Ohwi ex Karton. (Gesneriaceae), is described and illustrated. This species resembles the widespread Rhynchoglossum obliquum Blume. A new combination, Codonoboea kjellbergii (B.L.Burtt) Karton., is proposed to accommodate the former Henckelia kjellbergii B.L.Burtt.
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The genus Didymocarpus has for long been used as a dumping ground for species of different affinities and different geographical origin. Following recent work on Didymocarpus and the associated genera Loxo- carpus, Codonoboea, and Platyadenia, which resulted in a much narrower delimitation of Didymocarpus, the re-instatement and enormous expansion of Henckelia and the erection of the new genus Hovanella (WEBER & BURTT 1998d), a list is presented in which the current position of the species formerly referred to these genera can be found. Types are quoted for all species except for those belonging to Didymocarpus s.str., for which the quotations are given in WEBER & al. (2000). Notes and references to recent literature are added.
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Based on molecular data and morphology, Metabriggsia is reduced to synonymy with Hemiboea and its two species transferred to that genus.
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Gesneriaceae is the largest family of dicotyledonous herbs in Malaysia with 23 genera and about 400 species. The great majority grow in the ground layer of the rain forest (e.g., Cyrtandra and Henckelia species) or are epiphytes (e.g., Aeschynanthus) or climbers (e.g., Agalmyla), while others are mostly restricted to limestone (e.g., Chirita, Monophyllaea and Paraboea). Two life strategies are seen in the family: deep-shade rainforest species (e.g., Henckelia platypus) are slow-growing, live for up to 20 years but their seedlings are rarely seen. In contrast, limestone species produce copious seedlings, are fast-growing, early flowering and short-lived, most succumb to drought and are annuals, Monophyllaea horsfieldii is slightly longer lived but dies when its hypocotyl can no longer support the lamina. Common ways of extending the life span of the individual is the decumbent stem, or in monocarpic species by producing sprouts from the stem base, or rarely from leaf cuttings. Flowering patterns includes continuous, gregarious or seasonal flowering. Observations on pollination are lacking; flower visitors observed include birds for Aeschynanthus, bees for Henckelia species, butterflies for Monophyllaea. Self-pollination may occur in some species. Seed dispersal is probably by wind (e.g., Aeschynanthus and some Paraboea) and by ballistic force of water drops from the forest canopy (e.g., Epithema, Henckelia). None grows in secondary or disturbed habitats so the major threat to survival is habitat disturbance.
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Chirita D. Don, a large genus in the subfamily Cyrtandroideae of Gesneriaceae, has been the subject of much debate whether it is a natural group or not. In addition, the highly heterogeneous Chirita has also been very problematic with regard to delimitation and subdivision. Here we used the nrDNA internal transcribed spacer and cpDNA trnL-F for molecular phylogenetic analaysis, combined with morphological data. Our results suggest that Chirita is an artificial, polyphyletic genus. The most important character that defines Chirita, the dorso-ventrally oblique and bilamellar stigma, has evolved convergently in different clades of diandrous Cyrtandroideae. Chirita sensu strict only includes the species of Chirita sect. Chirita, whereas Chirita sect. Microchirita is an independent clade located at the basal node of the phylogenetic tree. Chirita sect. Liebigia is closely related to Didymocarpus with an entire stigma unlike other species of Chirita. The species of Chirita sect. Gibbosaccus, Chiritopsis, Primulina, and Wentsaiboea form a monophyletic group that is sister to a strongly supported clade comprising four monotypic genera Paralagarosolen, Calcareoboea, Petrocodon, and Tengia. We further analyzed the morphological evolution of Chirita and identified a series of morphological synapomorphies for the monophyletic groups revealed herein, and thereby provide a taxonomic treatment in this study.
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Of the 25 species presently named in Henckelia sect. Loxocarpus (Gesneriaceae), 10 occur in Peninsular Malaysia (with one species extending into Peninsular Thailand), 12 in Borneo, two in Sumatra and one in Lingga (Indonesia). One species from Peninsular Malaysia is excluded from the section. A key to species in Peninsular Malaysia is provided, together with descriptions and distributions. Three new species are described: Henckelia anthonysamyi Banka, H. sekayuensis Banka & Kiew and H. sericiflava Kiew & Banka. All Peninsular Malaysian species are endemic, many narrowly so – eight are presently known from just one or two localities.