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Camellia puhoatensis, a new species from Vietnam 1
Camellia puhoatensis (Sect. Archecamellia – Theaceae),
a new species from Vietnam
Danh-Hùng Nguyễn1, Văn-Dũng Lương2, i-Hương Lê3,
Quốc-ành Trần4, Ngọc-Đài Đỗ1,5, Ngọc-Sâm Lý1,6
1 Graduate University of Science and Technology, VAST, 18 Hoang Quoc Viet, Cau Giay District, Ha Noi,
Vietnam 2 Faculty of Biology, Da Lat University, 1 Phu Dong ien Vuong Road, District 8, Da Lat City,
Lam Dong Province, Vietnam 3 School of Natural Science Education, Vinh University, 182 Le Duan, Vinh
City, Nghe An Province, Vietnam 4 Department of Science and Technology Nghe An, 75, Nguyen i Minh
Khai, Vinh City, Nghe An Province, Vietnam 5 Faculty of Agriculture, Forestry and Fishery, Nghe An College
of Economics, 51 Ly Tu Trung, Vinh City, Nghe An Province, Vietnam 6 Department of Biological Resources,
Institute of Tropical Biology, VAST, 85 Tran Quoc Toan, District 3, Ho Chi Minh City,Vietnam
Corresponding author: Ngọc-Sâm Lý (lysamitb@gmail.com)
Academic editor: C. Morden |Received 15 December 2019| Accepted 8 June 2020 |Published 16 July2020
Citation: Nguyễn D-H, Lương V-D, Lê T-H, Trần Q-T, Đỗ N-D, Lý N-S (2020) Camellia puhoatensis (Sect. Arche-
camellia – eaceae), a new species from Vietnam. PhytoKeys 153: 1–11. https://doi.org/10.3897/phytokeys.153.49388
Abstract
A new species of eaceae, Camellia puhoatensis N.S. Lý, V.D. Luong, T.H. Le, D.H. Nguyen & N.D.
Do, sp. nov., is described and illustrated from Nghe An Province, North Central Coastal Region, Viet-
nam. It is most similar to C. chrysanthoides, C. avida and C. petelotii within sect. Archecamellia in shape
and colouration of leaf, petal, ovary and glabrous stamen, but diers by its young puberulous shoot,
mature leaf sparsely puberulous abaxially and leaf base rounded or broadly obtuse, petiole and pedicel
puberulous, tepals 12–13, ovary and style pubescent. e comparison between the new species and C.
velutina and C. dormoyana is presented. Data on distribution, ecology, phenology, use and provisional
conservation assessment of the new species are given along with an illustration and a colour plate.
Keywords
Camellia, section Archecamellia, taxonomy, eaceae, Vietnam
PhytoKeys 153: 1–11 (2020)
doi: 10.3897/phytokeys.153.49388
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Danh-Hùng Nguyễn et al. / PhytoKeys 153: 1–11 (2020)
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Introduction
Camellia Linnaeus (1753) is the largest genus of the family eaceae, with recent authors
recognising species between 120 (Ming and Bartholomew 2007) and 280 (Chang 1981;
Gao et al. 2005), distributed widely in East and Southeast Asia, from the Himalayas east
to Japan and Indonesia (Chang and Ren 1998; Ming and Bartholomew 2007). e high-
est species diversity is found in China and Vietnam (Chang and Ren 1998; Ming 2000;
Orel and Wilson 2012b). Camellia is distinguished from other genera of eaceae by its
usually large and apically dehiscent capsules and wingless (semi-)globose or polygonal
seeds with an umbilicate hilum (Ming and Bartholomew 2007). e general introduc-
tion to the genus, with particular focus on Vietnam, was given in recent publications by
various authors (e.g. Orel et al. 2012, 2013, 2014a, b; Luong et al. 2016a; Nguyen et al.
2018; Pham et al. 2019; Do et al. 2019a, b). So far, more than 75 species of Camellia have
been reported in Vietnam, with many localised endemic species (e.g. Pitard 1910; Gagne-
pain 1941; Rosmann 1999; Tran 1998a, b; Pham 2000; Hakoda and Tran 2001; Hakoda
et al. 2007; Orel 2006; Orel and Wilson 2010a, b, 2012a, b; Orel and Curry 2014; Orel
et al. 2012, 2013, 2014a, b; Tran et al. 2012; Tran and Luong 2012, 2013; Tran and Le
2013, 2015; Luu et al. 2015, 2018; Luong et al. 2016a, b; Le et al. 2017; Nguyen et al.
2018; Pham et al. 2019; Do et al. 2019a, b), but the actual number is expected to be
higher in the near future (Le and Luong 2016, Do et al. 2019b).
During recent extensive oristic surveys in the North Central coastal region in Vi-
etnam, several interesting species of Camellia in yellow ower were collected by one of
us (N.-D. Do) and colleagues in 2018–2019 (e.g. Tran and Luong 2013; Tran and Le
2015; Luong et al. 2016a, b; Le et al. 2017; Nguyen et al. 2018; Pham et al. 2019; Do
et al. 2019a, b). Critical examination of living owers, dried specimens and comparison
with type material and protologues of all related yellow Camellia in Vietnam and China
(e.g. Sealy 1958; Chang 1981; Chang and Bartholomew 1984; Gao et al. 2005; Ming
2000; Ming and Bartholomew 2007; Pham 2000; Orel and Curry 2014) led to the dis-
covery of several new taxa, two of which were recently described and named C. pukhan-
gensis D.N. Do, D.V. Luong, S. T. Hoang & H.T. Le and C. ngheanensis N.D. Do, V.D.
Luong, N.S. Ly, T.H. Le & D.H. Nguyen (Do et al. 2019a, b), while some other col-
lections are still awaiting description. In this paper, we describe a further new Camellia
from the Pu Hoat Nature Reserve, Nghe An Province, Vietnam. e overall plant habit,
somewhat ovate leaf blades, orbicular sepals and bright yellow tepal of these plants in Pu
Hoat NR show similarities with C. chrysanthoides H.T. Chang, C. avida H.T.Chang,
C. petelotii (Merr.) Sealy and C. dormoyana (Pierre) Sealy (Sealy 1949, 1958; Chang
1979, 1981). However, it shows signicant dierences in its vegetative and oral struc-
tures (see Table 1) and we describe it here as a new species to science, C. puhoatensis.
Materials and methods
e descriptions are mainly based on measurements from mature individuals of living
plants in the eld, supplemented by measurements from herbarium specimens. Type
Camellia puhoatensis, a new species from Vietnam 3
specimens of the most closely-related species of yellow camellias were examined from
the following herbaria: DLU, HN, P, NSW and VNM (herbarium codes follow iers
2018). Hi-resolution digital images available were also accessed from botanical websites
(e.g. https://science.mnhn.fr/, http://www.cvh.org.cn/, https://avh.ala.org.au/, https://
plants.jstor.org/). All morphological characters were described using the general termi-
nology and standard works of Sealy (1958), Chang (1981), Chang and Bartholomew
(1984), Chang and Ren (1998), Gao et al. (2005), Ming (2000) and Ming and Bar-
tholomew (2007). e conservation status was assessed, based on eld observations in
accordance with the IUCN Red List Categories and Criteria version 3.1 (IUCN 2017).
Taxonomic treatment
Camellia puhoatensis N.S. Lý, V.D. Luong, T.H. Le, D.H. Nguyen & N.D. Do, sp. nov.
urn:lsid:ipni.org:names:77210595-1
Figures 1, 2
Diagnosis. Camellia puhoatensis is morphologically similar to C. chrysanthoides, C. a-
vida and C. petelotii, but diers in having young puberulous shoots, mature leaves
sparsely puberulous abaxially with leaf bases rounded or broadly obtuse, petioles and
pedicels puberulous, tepals 12–13 and the ovary and styles pubescent.
Type. VIETNAM. Nghe An Province: Que Phong District, Dong Van Commune,
Pu Hoat NR, 19°43'31"N, 105°05'43"E, 270 m elev., 30 December 2018, Do Ngoc
Dai, Le i Huong, Nguyen Danh Hung, DHH-682 (holotype VNM; isotypes P, DLU).
Description. Shrub to tree, 2–7 m tall; young shoots puberulous then glabrous when
aging, purple towards terminals; semi-mature branches brown, smooth, glabrous, leaf
scars prominent; adult branches and trunk light grey, smooth with lighter-coloured
patches and covered by lichens; axillary leaf buds rudimentary, roughly triangular, at,
with rounded apex, pubescent, brown, bud scales small but prominent, 1–3 mm long.
Leaves: juvenile leaves forming a narrow tube when young, soft, somewhat pendulous,
purple in colour; young leaves slightly serrate, shiny, purple, adaxially glabrous, abaxi-
ally puberulous; developing leaves descending, narrow, shiny, purple to green-purple
tinted, abaxial surface puberulous; mature leaves serrate, irregularly towards the apex,
17–23 × 5.0–6.5 cm; petiole 8–16 × 4–5 mm, puberulous; lamina thin, coriaceous,
oblong ovate or oblong, leaf apex acuminate or narrowly acuminate, base rounded or
broadly obtuse, adaxially dark green and glabrous, abaxially pale green and sparsely
puberulous; primary vein continues as a shallow channel on the adaxial side of the
petiole, 2.0–2.5 mm wide proximally, less than 1.0 mm distally, proximally light green
and shiny on both sides; secondary venation pinnate, indistinctly brochidodromous,
partially eucamptodromous on some leaves, with 10–13 pairs; midribs and lateral
veins sunken adaxially; veins distinct proximally, less so towards the apex and the mar-
gins; tertiary venation very indistinct, sometimes lacking, more prominent at the leaf
margins. Flowers usually solitary, sometimes together in groups of 2 owers borne on a
short bracteate shoot, terminal, rarely axillary, lacking scent, 4.5–6.0 cm in diameter;
Danh-Hùng Nguyễn et al. / PhytoKeys 153: 1–11 (2020)
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Figure 1. Camellia puhoatensis. A Leaf, adaxial view B venation detail of leaf (abaxial surface) C ower
(lateral view) D ower (top view) E bracteoles (inner surfaces shown) F sepals (adaxial surfaces) G petals
(adaxial surfaces) H androecium (one part) I stamen J gynoecium (with sepals and petals). Drawn from
the holotype by Van-Dung Luong.
Camellia puhoatensis, a new species from Vietnam 5
pedicel stout, covered by purplish-red perulae, 7–10 mm long, puberulous; ower buds
unevenly globose in shape, 2.2–2.6 × 2.0–2.3 cm, yellowish-red tinted, open ow-
ers somewhat circular; bracteoles (sensu Sealy 1958) 3–4, opposite, orbicular, 1.5–2.5
× 1.5–3.0 mm, abaxially red to yellow-red tinted, adaxially paler, glabrous, margins
ciliate, persistent; sepals 5, persistent, orbicular or subglobose, 0.6–1.5 × 0.8–1.8 cm,
abaxially dull red and pubescent, adaxially pale yellowish and glabrous, margins cili-
ate; petals 12–13, arranged in 3 whorls, bright yellow, sometimes with large red patch
on the outer ones; outermost whorl comprising 3 or 4 petals, orbicular to broadly
obovate, 2.2–2.8 × 1.6–2.3 cm, abaxially pubescent, adaxially glabrous; middle whorl
comprising 4 or 5 petals, broadly obovate, 2.4–3.3 × 1.8–2.5 cm, abaxially pubescent,
adaxially glabrous; innermost whorl of 3 or 4 petals, orbicular to broadly obovate,
2.3–2.5 × 1.7–2.2 cm, abaxially pubescent, adaxially glabrous, basally united with out-
ermost laments 5–7 mm. Androecium numerous stamens, in 4–5 whorls, light yellow,
2.5–2.8 cm long, glabrous; outer laments basally united for 1.5–1.8cm forming a
cup, inner ones basally united for 3–5 mm, free above union; anthers yellow, 2.2–2.8 ×
1–1.5 mm, with two longitudinal striations, dorsixed. Gynoecium superior, 3–(4)-loc-
uled, ovoid terminating into 3–(4) styles, 2.5–3.0 × 3.0–3.5 mm, slightly longitudinal
striations, pubescent, 2 ovules per locule; styles free to the base, 1.8–2.3cm long, pu-
bescent. Capsule not seen.
Phenology. Flowering from November to January of the next year.
Distribution and habitat. Camellia puhoatensis is currently found only from the
type locality. It grows on moist fertile and sandy soils along mountain streams and hill
slopes in evergreen broad-leaved forests in Pu Hoat Nature Reserve, Vietnam, at eleva-
tions of 270–450 m.
Provisional conservation assessment. At present, only a single population of
about 300 scattered mature individuals of Camellia puhoatensis was observed in the
type locality, with an estimated extent of occurrence (EOO) less than 100 km2 and an
area of occupancy (AOO) less than 1 km2. e population is highly threatened due
to loss of habitat within its range and high market demands for wild, yellow-owered
camellias which are intensively collected for sale by local people. erefore, C. puhoat-
ensis is preliminarily categorised as Critically Endangered [B1ab (i, ii, iii) + 2ab (ii, iii),
D], according to the IUCN Categories and Criteria (IUCN 2017).
Etymology. e specify epithet ‘puhoatensis’ refers to the type locality.
Uses. Leaves and owers were harvested and used for tea by the local people.
Additional specimens examined. Paratypes. V. Nghe An Province: Que
Phong District, Dong Van Commune, Pù Hoạt NR, 19°48'45"N, 105°5'39"E, 320m
elev., 2 September 2018, Đỗ Ngọc Đài, Nguyễn Danh Hùng, Lê ị Hương, DHH 120
(VNM); the same locality, 19°48'31"N, 105°05'43"E, 280 m elev., 16 January 2019,
Đỗ Ngọc Đài, Nguyễn Danh Hùng, Lê ị Hương, DHH 790 (DLU), DHH 791 (HN).
Vernacular name. Vietnamese language: Trà hoa vàng pù hoạt.
Taxonomic notes. e current infrageneric classication of Camellia is derived
from three previous publications (Sealy 1958; Chang and Bartholomew 1984; Ming
2000) and was based on the assessment of morphological characteristics. e taxo-
Danh-Hùng Nguyễn et al. / PhytoKeys 153: 1–11 (2020)
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Figure 2. Camellia puhoatensis. A young shoot B terminal buds C solitary bud and axillary ower (side
view) D ower and pollinated ower (side view) E close-up of ower (front view) F immature fruit G a
part of branch showing leaves abaxial and opening ower H leaves adaxially I bracteoles J sepals K petals
L androecium with stamens M gynoecium (with sepals and styles). Photos by Ngoc-Dai Do, the colour
plate prepared by Ngoc-Sam Ly.
Camellia puhoatensis, a new species from Vietnam 7
nomic system of Sealy (1958) and Chang and Bartholomew (1984) are used to de-
scribe and determine the placement of new taxa within Camellia. ese systems are the
most detailed and comprehensive study of the genus and also provide the basis for our
current understanding of the genus. e taxonomic system of Ming (sensu Ming and
Bartholomew 2007) was used for supplementary data only as it appears to be super-
cially similar to the system of Sealy (Orel and Wilson 2010a). In this paper, we have
followed the Sealy’s (1958) taxonomic system to consider the placement of the new
species within Section Archecamellia Sealy of Camellia. Morphologically, C. puhoatensis
possesses a solitary or paired owers at terminal (sometimes axillary), stout, thick and
erected pedicel, 3–4 persistent bracteoles, 5 persistent sepals (undierentiated bracte-
oles and sepals), large yellow owers with 12 tepals that are inner ones basally connate
and adnate to androecium, androecium free above the union with the petals or unied
for some distance to form a eshy cup, laments glabrous, gynoecium 3(–4)-loculi,
styles 3(–4) and free to the base. ese characteristics are not only identical to the spe-
cies of sect. Archaecamellia Sealy (sensu Sealy 1958; sensu Chang and Bartholomew
1984), but also share with species of sect. Stereocarpus which possesses 2 or 4 bracteoles
(sensu Sealy 1958), terminal owers (sensu Chang and Bartholomew 1984), stamens
in 3–4 whorls, ovary with 3–5 locules (sensu Ming 2000), as well as sharing with spe-
cies of sect. Chrysantha Chang, such as distinct peduncle, small oral bracteolates,
yellow owers, three carpels of gynoecium and separate styles (sensu Chang 1979).
As characterized by Sealy (1958), sect. Archaecamellia shares several traits with sect.
Stereocarpus. ese include a solitary and erected ower at the end of the branches,
persistent bracteoles and sepals, stamens united with the petals and glabrous laments.
However, traits that are distinctive to sect. Archaecamellia include (6–)11–16 indistinct
bracteoles and sepals, 8–14 petals, glabrous or pubescent gynoecium and 3 or 5 free
styles (vs. 2 or 4 bracteoles and 5 or 6 sepals, ca. 12 petals, glabrous gynoecium and
a single style in sect. Stereocarpus). e sect. Chrysantha also shares several traits with
sect. Archaecamellia in having yellow and pedicellate owers, persistent bracteoles and
sepals, glabrous or hairy laments and gynoecium, but it can be distinguished from
sect. Archaecamellia by the axillary owers, distinct bracts and sepals and 3–5 cleft
styles (Chang 1979). Section Archaecamellia is currently comprised of 19 species (Sealy
1958; Ming 2000; Orel and Wilson 2012a; Do et al. 2019a). e new species is most
similar to C. chrysanthoides H.T.Chang, C. avida H.T.Chang and C. petelotii (Merr.)
Sealy in having the same plant habit, somewhat oblong leaves, yellow owers, glabrous
3-loculed gynoecium with 3 styles free to the base. A detailed morphological compari-
son between C. puhoatensis and these three species is provided in the above diagnosis
and in Table 1. Moreover, C. puhoatensis also resembles C. dormoyana (Pierre) Sealy
of sect. Stereocarpus (Sealy 1958) and C. velutina V.T. Pham et al. of sect. Chrysantha
(Pham et al. 2019) by somewhat oblong leaves, yellow owers and glabrous stamens.
However, Camellia dormoyana is easily distinguished from C. puhoatensis by having
the young shoots, mature leaves and petioles all glabrous, the sessile pedicel and 5–6
bracteoles abaxially velutinous, the abaxial petals silky velutinous, the ovary being gla-
brous and with ve locules and the styles united for their entire length and glabrous.
Danh-Hùng Nguyễn et al. / PhytoKeys 153: 1–11 (2020)
8
Similarly, C. velutina is readily distinguished from C. puhoatensis by its glabrous young
shoots, mature leaves and petioles, sepals that are silky velutinous abaxially and velu-
tinous adaxially, the 10 (occasionally 11) petals that are silky velutinous and glabrous
ovary and style (see Table 1).
Acknowledgements
We are grateful to the manager and sta of Pu Hoat Nature Reserve and to Mr. Tran
Quoc anh, director of Nghe An Provincial Department of Science and Technology
for their helpful cooperation during the eldwork. We thank Mr. Bruce Maslin (PERTH
herbarium, Australia) for helping to improve the English text and to the anonymous
reviewers for their constructive comments that helped improve the manuscript.
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