PhytoKeys 34: 33–46 (2014)
Lithocarpus orbicarpus (Fagaceae), a new species of Stone Oak...
doi: 10.3897/phytokeys.34.6429
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Lithocarpus orbicarpus (Fagaceae), a new species of
Stone Oak from Phang Nga province,Thailand
Joeri S. Strijk1, Sukontip Sirimongkol2, Sukid Rueangruea2,
Nikom Ritphet2, Voradol Chamchumroon2
1 Ecological Evolution Group, Key Laboratory of Tropical Forest Ecology, Xishuangbanna Tropical Botanical
Garden, Chinese Academy of Sciences, Menglun, Mengla, Yunnan, 666303, PR China 2 Forest Herbarium
(BKF), Department of National Parks, Wildlife and Plant Conservation, 61 Phahonyothin Rd., Chatuchak,
Bangkok 10900, Thailand
Corresponding author: Joeri S. Strijk (joeristrijk@xtbg.org.cn)
Academic editor: H. Schaefer | Received 14 October 2013 | Accepted 2 February 2014 | Published 11 February 2014
Citation: Strijk JS, Sirimongkol S, Rueangruea S, Ritphet N, ChamchumroonV (2013) Lithocarpus orbicarpus (Fagaceae),
a new species of Stone Oak from Phang Nga province, Thailand. PhytoKeys 34: 33–45. doi: 10.3897/phytokeys.34.6429
Abstract
A new species is described, Lithocarpus orbicarpus Strijk, collected from Ton Pariwat Wildlife Sanctuary,
Mueang district, Phang Nga Province in the Peninsular Floristic Region of Thailand. We provide the first
technical illustrations and colour photographs of the new species, as well as a description of its conservation status and the collecting locality. The species can be easily distinguished by its unique orbicular
acorns, each covered with a dense pattern of irregularly placed scales, which completely conceal the nut,
except for a tiny apical pore, and which are arranged in a dense cluster on an erect woody spike. We also
provide an amendment to the existing diagnostic key to Lithocarpus, and discuss important differences
with morphologically similar species found in Thailand and the surrounding region.
Keywords
Lithocarpus orbicarpus, new species, Stone Oaks, Fagaceae, Thailand, Ton Pariwat Wildlife Sanctuary
Introduction
Lithocarpus Blume is the second largest genus within Fagaceae, with over 300 species described (Camus 1948, 1954; Phengklai 2008; Soepadmo 1972; Wu et al.
1999). Members of the genus, commonly called Stone Oaks, can be found in (sub-)
Copyright Joeri S. Strijk et al. This is an open access article distributed under the terms of the Creative Commons Attribution International License
(CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
34
Joeri S. Strijk et al. / PhytoKeys 34: 33–45 (2014)
tropical broad-leaved evergreen forests near sea level to mixed forests at altitudes of
over 3200 m. The geographic distribution of Lithocarpus roughly covers southern
and south-eastern Asia from eastern India to southern Japan, to the Philippines and
southward to New Guinea. Lithocarpus densiflorus (Hook. & Arn.) Rehder, the only
North American member of the genus was recently moved to a new monotypic genus (Manos et al. 2008). All species within Lithocarpus are trees, ranging from small
understory elements, to very large emergent trees. Many species have a geographically
wide distribution and in some locations may constitute the main canopy forming elements together with other Fagaceae (Lithocarpus, Quercus, Castanopsis), Lauraceae and
Theaceae. Leaves are simple, entire, rarely serrate, generally glabrous and mostly spirally arranged. Male and female flowers are white to pale whitish-yellow. Genders can be
either on separate inflorescences, on the same inflorescence mixed throughout, or with
female flowers below and male flowers terminal, arranged with dense indumentum on
erect spikes. Male flowers are solitary or in clusters of three or more, with the perianth
campanulate or cup-shaped, usually 6-lobed, partially united; stamens generally 12.
Female flowers solitary or in clusters of three, perianth like male flowers but smaller;
12 staminodes; 3(-4) styles (Camus 1948, 1954; Phengklai 2008).
Previous studies and the most recent treatment for Thailand have recovered 57
species of Lithocarpus (Barnett 1940; Phengklai 2004, 2008). During fieldwork in
remote Ton Pariwat Wildlife Sanctuary, carried out as part of ongoing research on
the genomics, systematics, biogeography and evolution of Asian Fagaceae, we made
collections of an individual tree with unique features that could not be matched with
any previously described taxa in Fagaceae. After careful examination of herbaria and
literature, comparison of other specimens collected during the fieldwork and consultation of specialists on the regional flora, we report this collection here as a new species,
placed within the genus Lithocarpus.
Taxonomy
Lithocarpus orbicarpus Strijk, sp. nov.
urn:lsid:ipni.org:names:77135982-1
http://species-id.net/wiki/Lithocarpus_orbicarpus
Fig. 1, 2
Type. THAILAND, Ton Pariwat Wildlife Sanctuary, Mueang district, Phang Nga
Province, 8°37'25"N; 98°33'14"E; alt. 455 m, 16 July 2013, Chamchumroon et al.
5823 (Holotype: BKF; Isotypes: E, K, L, SING).
Diagnosis. Lithocarpus orbicarpus is a small-medium sized tree. It differs from similar species by its unique orbicular acorns, each covered with a dense pattern of irregularly placed scales, which completely conceal the nut, except for a tiny apical pore, and
which are arranged in a dense cluster on an erect woody spike. Unique for Thai species
of Lithocarpus, almost the entire surface of the round nut is covered with scar area (re-
Lithocarpus orbicarpus (Fagaceae), a new species of Stone Oak...
35
ceptacle tissue), leaving only the topmost part of the nut covered with a thin vestigial
exocarp layer. Pending discovery of additional individuals, the species appears to be locally restricted to low-mid-elevation forests in the peninsular region of Thailand.
Description. Small-medium sized tree, up to 15 m tall. Bark smooth to slightly
rough grey-green, with superficial horizontal lines. Sapwood white to yellow, with
inner bark ridges forming light brown longitudinal slits in sapwood surface. Branches
dark brown to grey brown, mostly glabrous, densely lenticellate; young twigs, leaf buds
and old fruits with short, soft (occasionally long) gray indumentum. Leaf buds tiny
and terminal buds solitary. Leaves simple; lamina elliptic to oblanceolate with (strongly) acuminate tip, 11.0–22.3 × 4.4–7.1 cm. Margin entire. Leaves often with slightly
asymmetric lamina. Leaf apex acuminate to strongly acuminate, leaf base cuneate to
slightly attenuate. Both surfaces generally glabrous except emerging leaf buds, terminal
shoots and young leaves, which have soft grey indumentum. Young leaves light green,
but turning dark green above and glaucous below when older. Venation. Pinnately
veined; secondary venation discretely anastomosing near the leaf margin. Pairs of secondary veins 9–13, slightly raised and clearly visible on underside of leaf. Peduncles
carrying fruits 5–11 cm long, up to 1 cm thick at the base, glabrescent, grey-brown
and densely lenticellate. Male and female inflorescences not seen. Infructescence a woody
spike, terminal, up to 15–21 cm long. Fruits sessile on thick woody peduncle, closely
pressed against each other, but walls of individual units not fused. Number of fruits
per infructescence very variable, ranging from 9–20 units. Acorn. Orbicular, globose,
2.7–3.4 by 2.9–3.5 cm (including cupule) and covered with glabrous, semi-concentric
interlocking ridges when young, which transform over time into ridges with irregularly
placed scales. Cupule nearly completely enclosing the nut, indehiscent, but showing
small cracks when mature; fruit wall up to 4–6 mm thick, apical pore very small, 1–4
mm wide, exposing the persistent punctiform styles (3) and a tiny fraction of vestigial
exocarp. Young cupule walls light green, ridges light to dark brown. Old cupule walls
turning light brown to yellow-brown and pubescent with short (occasionally long),
greyish-yellow indumentum. Nut 1 in each cupule, ball shaped, globose, 2.4–2.9 by
2.6–3.0 cm. Up to 95% of surface area of the nut made up of scar area (receptacle
tissue), upper 5% of surface area of the nut slightly raised and made up of vestigial
exocarp layer. Nut scar pale yellow-whitish, tiny exocarp layer light brown. Scar area
covered with deep groves and red-brown to purplish vein-like lines, stretching down to
the base of the nut. Up to 5/6 of the scar area of the young nut (from the base upward)
covered with dotted pattern of small depressions. Cotyledons black when dried.
Flowering and fruiting. Flowering unknown, but thought to be around November-January. Infructescences on the ground in July, fruiting commencing possibly as
early as April-May.
Distribution. This species is only known from Thailand, and has not been recorded outside Ton Pariwat Wildlife Sanctuary, Mueang district, Phang Nga Province.
During our field survey, we found only one individual tree, located on a gentle sloping
section of closed dense forest. Additional survey work will have to be undertaken to
determine the actual population size of this species within the wildlife sanctuary.
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Joeri S. Strijk et al. / PhytoKeys 34: 33–45 (2014)
Figure 1. Lithocarpus orbicarpus Strijk, sp. nov. Chamchumroon et al. 5823 (BKF). Technical illustration.
A Habit B Detail of glabrous young fruit with ridges and apical pore C Detail of interior of young fruit,
showing nearly complete fruit scar, covered umbo and ‘pitted pattern’ on the nut surface D Infructescence
with ripe fruits showing highly irregular scaly patterns on the fruit exterior E Details of seed, from left to
right: bottom view, side view, top view. Note venation and crevice pattern on surface of fruit, and cover of
the umbo section. All drawings by J.S. Strijk.
Lithocarpus orbicarpus (Fagaceae), a new species of Stone Oak...
37
Ecology. This species grows in dense hillside forest at an elevation of around 450 m.
Vernacular name. Unknown.
Etymology. Lithocarpus orbicarpus is named after its unique orbicular acorns, arranged densely clustered on an erect spike, of which the cup almost completely encloses the nut (except for a tiny apical pore). The exterior of the cup is covered with a dense
pattern of ridges, transforming with age into horizontal and vertical lines with irregularly placed scales. Apart for a tiny portion of the umbo which is covered with vestigial
exocarp, the exterior surface of the nut is completely covered by scar. Although the
cupule nearly completely covers the nut, the latter is free and not fused to the wall of
the cupule – a condition which occurs throughout the genus (Cannon 2001; Cannon
and Manos 2001). Upon drying, the shrinking of the cotyledons inside the nut causes
the whole infructescence to make a rattling sound when shaken.
Discussion. Thailand has a total of 121 species, 2 subspecies and 2 varieties of
Fagaceae in four genera indigenous to the country. Of these, nine species (Castanopsis: 4; Quercus: 1; Lithocarpus: 4) and 1 subspecies (Quercus: 1) are national endemics
(Bunpha et al. 2011; Phengklai 2008). Three of the endemic Lithocarpus species (including L. orbicarpus) are restricted to small ranges in the peninsular region. Of the 58
Lithocarpus known to occur in Thailand (including this newly described species), 43
species (74%) can be found in the Peninsular floristic province making it the most diverse floristic region (in terms of Lithocarpus species) followed by the North (25 species
– 43%). While the Northern (and North-eastern) floristic province is characterized by
species that reach high elevation habitat (>75% can be found between 1200–2500 m),
the Peninsular complement includes species that are restricted to lower elevations (21
species restricted <500 m asl). With two exceptions (L. falconeri (Kurz) Rehder and
L. erythrocarpus (Ridl.) A.Camus), none of these species are shared with the North (or
North-east). In contrast, four of the Peninsular lowland species (L. lucidus (Roxb.)
Rehder; L. maingayi (Benth.) Rehder; L. reinwardtii (Korth.) A.Camus; L. tubulosus
(Hickel & A.Camus) A.Camus) are shared with the South-eastern floristic province,
in addition to L. elephantum (Hance) A.Camus and L. pierrei (Hickel & A.Camus)
A.Camus, which can only be found in the South-eastern region.
Lithocarpus in Thailand can be further characterized based on their acorn
(cupule+nut) properties. The open- or semi-open type, in which the cupule shape ranges from saucer shaped-flat to covering up to 4/5th of the nut is present in 37 species
(64%), while the remaining 18 species have cupules that entirely cover the nut, in most
cases leaving a tiny portion of the umbo uncovered. With one exception (L. truncates
(King ex Hook.f.) Rehder), all closed-type species occur in the Peninsula, and eight are
restricted to it (two shared with the Southeast). L. orbicarpus resembles species in this
group, with its indehiscent and near-closed cupules, restricted geographical distribution
and presence in lower elevation habitat, but is clearly distinct from them based on the
structure and shape of the fruit, the infructescence and the properties of the nut.
During our field survey, we encountered and collected additional Fagaceae species, e.g.
Lithocarpus reinwardtii (Korth.) A.Camus (Burma, Cambodia, Malaysia, Indonesia), L.
sundaicus (Blume) Rehder (Malaysia, Indonesia, Brunei), L. cantleyanus (King ex Hook.f.)
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Joeri S. Strijk et al. / PhytoKeys 34: 33–45 (2014)
Figure 2. Lithocarpus orbicarpus Strijk, sp. nov. Chamchumroon et al. 5823 (BKF). Pictures from field
collection. A Branch with leaves B Young infructescences C Bark and sapwood D Fresh fruit – top view
E Fresh fruit – side view F Fresh fruit – bottom view G Dried fruit – top view H Dried fruit – side view
J Dried fruit – bottom view (G–J: blue lines demarcate vestigial exocarp; red lines demarcate scar area
(receptacle tissue)) K Dried infructescence L Young acorn, opened up to show dotted pattern of small
depressions and surface structure of the umbo M Cross-section of fried nut, showing seed coat and black
cotyledons N Terminal leaf, twig and very small terminal bud, showing grey indumentum O Young
emerging leaf with soft grey indumentum. All pictures by S. Sirimongkol and J. S. Strijk.
Lithocarpus orbicarpus (Fagaceae), a new species of Stone Oak...
39
Rehder (Burma, Malaysia, Singapore) and several, as of yet, unidentified collections. Additional species encountered in previous surveys in the wildlife sanctuary are: Castanopsis
javanica (Blume) A.DC. (Vietnam, Malaysia, Singapore, Indonesia); Castanopsis nephelioides King ex Hook. f. (Malaysia, Singapore); Castanopsis purpurea Barnett (Thailand (endemic)); Castanopsis wallichii King ex Hook. f. (Malaysia, Singapore, Indonesia); Castanopsis inermis (Lindl.) Benth. & Hook. f. (Burma, Malaysia, Singapore, Indonesia, Philippines); Lithocarpus bennettii (Miq.) Rehder (Malaysia, Singapore, Indonesia); Lithocarpus
clementianus (King) A.Camus (Malaysia, Indonesia); Lithocarpus eucalyptifolia (Hickel et
A.Camus) A.Camus (Burma, Vietnam, Cambodia); Lithocarpus falconeri (Kurz) Rehder
(Burma, Malaysia); Lithocarpus garrettianus (Craib) A.Camus (China, Burma, Laos, Vietnam); Lithocarpus lucidus (Roxb.) Rehder (India, Malaysia, Singapore, Indonesia, Brunei);
Lithocarpus macphailii (Henders.) Barnett (Malaysia, Indonesia); Lithocarpus wrayi (King)
A.Camus (Vietnam, Malaysia, Indonesia) and Quercus oidocarpa Korth. (Burma, Vietnam,
Malaysia, Indonesia). Out of these 17 species, at least 14 have a geographic distribution
that is primarily or completely located in the Sundaland biogeographic region. Conversely,
only three species have a distribution that is mostly or entirely contained within the Indochinese biogeographic region. Despite the paucity of detailed biological records on this
remote area, the distributional data shows us that the Fagaceae flora in Ton Pariwat Wildlife Sanctuary is distinctly Sundaic in composition, and differs substantially from forests in
the more northern Indochinese region. This is in fact not surprising, as the Ton Pariwat
Wildlife Sanctuary is located near the southern edge of one of the world’s major phytogeographic and zoogeographic transition zones: the ~500km stretch between the biogeographically distinct and well-known Isthmus of Kra – the narrowest part of the connection
between mainland Southeast Asia and the Malay Peninsula (10°30'N) – and the line between Kangar (Malaysia) - 6.°51’N, and Pattani (Thailand) 6°87'N. Within this relatively
short distance, an abrupt and major shift takes place from northern Indochinese flora and
fauna, to those distinct for the southern Sundaland region (Hughes et al. 2003; Meijaard
2009; Parnell 2013; Van Steenis 1950; Woodruff 2003; Woodruff and Turner 2009).
When looking at the Fagaceae flora in the wider region, some properties of L. orbicarpus resemble species in Peninsular Malaysia, such as Castanopsis nephelioides King
ex Hook.f., but can easily be distinguished from species within Castanopsis (D.Don)
Spach, based on the presence of bark ridges that penetrate the sapwood, the nut which
is not fused to the cup and the orbicular shape of the fruit. Congeneric species from the
Indo-Chinese zone that share some characters with L. orbicarpus are L. rouletii (Hickel
& A.Camus) A.Camus (but fruit with basal scar, flattened, dehiscent; South Vietnam);
L. pachycarpus (Hickel & A.Camus) A.Camus (but leaves with yellow indumentum,
cupules pear-shaped, acorns not orbicular; Vietnam-Laos); L. kontumensis A.Camus
(but cupules truncate, higher than acorn, cupules sometimes fused; Vietnam-Laos);
L. lepidocarpus (Hayata) Hayata (but cupules truncate, sometimes fused, fruit not orbicular; central and south Taiwan); and L. laoticus (Hickel & A.Camus) A.Camus (but
cupule ovoid, high elevation habitat (Tibet, south and central China, Vietnam) (Wu
et al. 1999). Within Thailand, L. orbicarpus is unique in its combination of properties,
and we outline some of the defining differences with Thai species in Table 1 below.
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Joeri S. Strijk et al. / PhytoKeys 34: 33–45 (2014)
Table 1. Morphological differences between L. orbicarpus and other Thai species of Fagaceae.
L. encleisocarpus
L. wrayi (King)
C. nephelioides King
A.Camus
A.Camus
ex Hook.f.
1. Nut wall
Free from the cup
Free from the cup
Free from the cup
Fused to the cup
Almost complete,
Almost complete, but small but raised umbo free Almost complete, but
2. Cupule
Enclosure complete.
apical pore showing flat umbo (±5-10%). Easily raised umbo free (±5enclosure
Indehiscent.
remains (≤5%). Indehiscent. dehiscent in irregular 15%). Indehiscent.
parts.
Ovoid, usually
3. Nut shape
Orbicular.
Ovoid to globose.
Broadly conical.
depressed to one
longitudinal side.
Spines absent. Small, flattened
Sparsely covered
scales present. Irregularly
Spines and scales
Alternate pseudowith short, woody
4. Cup
intersecting lines present. absent. Wall smooth, spines and free scales
spines, 2-3 branched
densely greenish- present; pseudo-spines
surface
Old acorns pubescent with
reclining and
short (occasionally long),
brown hairy.
incurved or erect.
decurved.
greyish-yellow indumentum.
Orbicular, symmetric; young
Obovoid, always
5. Acorn
fruits occasionally slightly
Ovoid or turbinate.
Broadly ovoid.
asymmetric, usually
shape
skewered in young and dense
flattened adaxially.
infructescences
Entire or serrate in the
6. Leaf
Entire throughout.
Entire throughout. Entire throughout.
upper half.
margin
7. Scar
Orbicular, covering ≥95%
Basal, slightly
Basal, concave, ca. 1.5
- (nut fused to wall).
position,
of the fruit, from the base concave, ca. 1 cm in
cm in diameter.
shape and size
upward.
diameter.
Sparsely sericeous
8. Nut
Glabrous.
Greyish pubescent.
- (nut fused to wall).
indumentum
then dull brown.
Characters
L. orbicarpus Strijk
This species is endemic to Thailand and is currently only known from one location in Ton Pariwat Wildlife Sanctuary. The sanctuary covers a region of low-lying
forested mountains with a total area of approximately 100,000 ha at the southern end
of the Phuket mountain range. As such it is an integrated part of the Southern Forest Complex of Thailand. The sanctuary is popular for its rich bird- and wildlife (e.g.
Blue-banded Kingfisher (Alcedo euryzona Temminck, Alcedinidae) and Whitehanded
Gibbons (Hylobates lar L., Hylobatidae) as well as rare flora, such as Rafflesia kerrii
Meijer (Rafflesiaceae). Its unique species composition, high diversity and relatively
intact forest structure underscore the importance of strengthening ongoing and future
conservation measures at Ton Pariwat Wildlife Sanctuary, as a key element of wider
conservation efforts in southern Thailand.
Updated key for the species of Lithocarpus occurring in Thailand
Following the treatment of Fagaceae for the Flora of Thailand (Phengklai 2008), no
further updates have been published. In the updated key we include here, we incorporate the identification of L. orbicarpus and add several additional corrections.
Lithocarpus orbicarpus (Fagaceae), a new species of Stone Oak...
41
Key to the thai species of Lithocarpus
(based on vegetative characters and acorns)
1
2
3
4
5
5
4
6
6
3
7
7
2
8
9
10
10
9
11
12
12
11
8
13
14
14
Outer surface of cupules with annular or lamellate markings or markings
lacking
Cupules without lamellae, chartaceous or subcoriaceous, enclosing nearly all
of the nut, more or less dehiscent when mature
Cupules weakly dehiscent from the apex, cupule surface distinctly undulate
with vertical and horizontal lines
Cupule urn-shaped
Cupule base broadly conical, much broader than apex, skin distinct with
many vertical filiform lines or without. Nut conical.............5. L. blumeanus
Cupule base obconic, much narrow than apex, surface distinct with 3-4 horizontal filiform lines. Nut obconical...................................... 33. L. maingayi
Cupule top or globe shaped
Cupule top-shaped, enclosing 4/5 of nut, surface with 2–6 distinct horizontal, filiform lines................................................................30. L. macphailii
Cupule globe-shaped, enclosing nut completely, except for a tiny section at
the apex, surface with distinct irregularly placed scales along 5–9 horizontal
and vertical lines................................................................ 36. L. orbicarpus
Cupules readily dehiscent into irregular parts from the top, surface with 2–5
filiform, undulate, horizontal lines
Cupules with 2 or 3 such lines..................................... 18. L. encleisocarpus
Cupules with 4 or 5 such lines.........................................37. L. pattaniensis
Cupules with distinct lamellae, coriaceous, enclosing a variable amount of the
nut, indehiscent
Cupule enclosing not less than 1/2 of the nut
Cupule enclosing about 1/2 of the nut
Nuts ovoid to conical at apex, scar shallowly concave or flattened..................
................................................................................................24. L. gracilis
Nuts subhemispheric or depressed at apex, scar deeply concave.....................
........................................................................................ 8. L. clementianus
Cupule enclosing not less than 3/4 of the nut
Cupules obconic, enclosing nut almost completely except around the umbonate apex
Nut longer than broad, ca. 1 by 0.7 cm..................... 26. L. hendersonianus
Nut shorter than broad, 1–2.7 by 2–3 cm......................... 32. L. magnificus
Cupules saucer-shaped, enclosing ca. 3/4 of the nut.............1. L. aggregatus
Cupule enclosing not more than 1/4 of the nut
Nuts hemispheric or depressed on both sides
Cupule enclosing 1/5 to 1/4 of the nut.............................39. L. platycarpus
Cupule enclosed only the base of the nut
42
15
15
13
16
17
17
16
18
18
19
19
1
20
21
22
23
23
24
24
22
25
25
26
26
21
27
28
28
27
29
30
Joeri S. Strijk et al. / PhytoKeys 34: 33–45 (2014)
Acorns sessile. Scar deeply concave..........................................15. L. eichleri
Acorns with stalk up to 0.5 cm long. Scar slightly concave........6. L. cantleyanus
Nuts conical to broadly ovoid, or with a dome-shaped apex
Cupule enclosing only the base of the nut
Acorns sessile. Leaves oblanceolate...........................................29. L. lucidus
Acorns with fruit-stalk up to 0.5 cm long. Leaves oblong...... 43. L. reinwardtii
Cupule enclosing ca. 1/4 of the nut
Nut with one horizontal ring around equator. Leaves ensiform to linearlanceolate.............................................................................. 28. L. loratefolius
Nut without horizontal ring. Leaves ovate, ovate-oblong or narrowly elliptical
Nut ovoid or conical. Cupules cup or saucer-shaped. Leaves ovate or ovateoblong, apex caudate.............................................................. 3. L. bancanus
Nut broadly ovoid. Cupules slightly obconical to saucer-shaped. Leaves narrowly elliptical............................................................................ 41. L. rassa
Outer surface of cupules with alternate lamellae (resembling fish scales) or
pseudospines
Mature cupules of one infructescence more or less fused together
Acorns broader than long, depressed both on top and at base. Cupules sauceror cupshaped or obconic, some hardly distinct from each other through fusion
Infructescences with densely arranged cupules
Cupules barely distinct, resembling a large gall.............. 13. L. echinophorus
Cupules distinct, saucer-shaped
Nut flattened or apiculate at apex, to 2.2 cm diam. Leaves cuneate at base....
................................................................................................ 16. L. elegans
Nut retuse at apex, not less than 3 cm diam. Leaves auriculate at base...........
........................................................................................... 2. L. auriculatus
Infructescences with spaces between cupules
Rachis of infructescence always with sub-branches. Acorns stalked................
....................................................................................... 34. L. mekongensis
Rachis of infructescence without sub-branches
Acorns sessile.............................................................................24. L. finetii
Acorns stalked..................................................................50. L. tenuinervis*
Acorns longer than broad, conical, ovoid or turbinate. Cupules cup-shaped
or cylindric
Rachis of infructescence always with sub-branches. Acorns stalked, nuts shining
Acorn up to 1 cm high. Rachis up to 4 mm in diam................7. L. ceriferus
Acorn not less than 1 cm high (to 2.5 cm). Rachis not less than 4 mm in
diam............................................................................... 40. L. polystachyus
Rachis of infructescence without sub-branches. Acorns sessile, nuts more or
less shining
Twigs glabrous or sparsely pubescent then glabrous
Cupules cup-shaped, enclosing up to 1/2 of the nut............12. L. dealbatus
Lithocarpus orbicarpus (Fagaceae), a new species of Stone Oak...
30
29
31
31
20
32
33
34
35
35
34
36
37
37
36
38
38
33
39
40
40
41
41
39
42
43
43
42
44
45
43
Cupules turbinate, enclosing the whole nut, open only around umbo...........
............................................................................................ 53. L. truncatus
Twigs ferruginous or tomentose
Leaves glabrous except along midrib. Cupules enclosing up to 1/3 of the
nut...............................................................................25. L. harmandianus
Leaves densely tomentose especially on lower surface. Cupules enclosing 1/2
of the nut..........................................................................27. L. lindleyanus
Mature cupules of one infructescence, free, not fused
Acorn longer than broad, conical, ovoid or obconical. Cupules cup- or saucer-shaped or obconic
Cupules enclosing nut completely or 2/3 of the nut
Cupules enclosing ca. 2/3 of the nut
Cupules slightly obconical-shaped, nuts hairy at style apex (if persistent).......
............................................................................................. 45. L. rufescens
Cupules cup or saucer-shaped.................................................. 16. L. elegans
Cupules enclosing nut completely, or up to the apex of the nut
Cupules dehiscent, obconic or ovoid
Cupules obovoid, sessile, surface with dense, long and narrow recurved pseudospines............................................................................. 42. L. recurvatus
Cupules ovoid, fruit stalk 2–3 mm long, surface finely ornamented with thin,
triangular lamellae throughout.....................................35. L. neo-robinsonii
Cupules indehiscent, ovoid, surface clothed with dense, triangular lamellae
Infructescences up to 18 cm long. Leaves up to 16 cm long...... 9. L. craibianus
Infructescences not less than 20 cm long. Leaves not less than 20 cm long....
..................................................................................... 19. L. erythrocarpus
Cupules enclosing up to 1/2 of the nut
Acorns stalked
Cupules slightly obconic. Leaves ovate, ovate-oblong or obovate...................
..........................................................................................48. L. sootepensis
Cupules cup-shaped or saucer-shaped
Cupules cup-shaped. Leaves lanceolate to lanceolate oblong.........47. L. siamensis
Cupules saucer-shaped to flattened. Leaves oblong to oblong-lanceolate........
...............................................................................................10. L. curtissii
Acorns sessile
Acorns (mature) not less than 3.5 by 2.2 cm
Cupule lamellae bearing pseudo-spined reflexed towards the base. Leaves
acute to obtuse at apex......................................................46. L. scortechinii
Lamellae curved towards the cupule apex. Leaves acuminate at apex..............
....................................................................................20. L. eucalyptifolius
Acorns (mature) up to 3 by 2.2 cm
Infructescence with acorns in clusters, but not fused
Nuts ovoid. Leaves usually curved to one side.................54. L. wallichianus
44
45
44
46
46
32
47
48
49
49
50
50
48
51
51
52
52
47
53
54
54
53
55
56
56
55
57
58
59
Joeri S. Strijk et al. / PhytoKeys 34: 33–45 (2014)
Nuts strongly apically depressed, occasionally conic. Leaves not curved.........
........................................................................................ 51. L. thomsonii
Infructescence with acorns solitary, with spaces between them
Cupules saucer or cup-shaped, limb recurved. Leaves not less than 12 cm
long...................................................................................... 21. L. falconeri
Cupules obconical, limb not recurved. Leaves up to 11 cm long................
...............................................................................................4. L. bennettii
Acorns broader than long, hemisphaeric-depressed
Cupules enclosing the nut completely or up to the apex of the nut
Cupules more or less up to the apex of the nut, lamellae with erect or reflexed
pseudospines which are not fused
Pseudo-spines erect or spreading. Leaves oblanceolate. Scar nearly 1/2 of the
nut.........................................................................................14. L. echinops
Pseudo-spines reflexed. Leaves oblong or oblanceolate
Infructescence with acorns packed close together, but not fused. Leaves
slightly cuneate at base.....................................................23. L. garrettianus
Infructescence with acorns solitary, with spaces between them. Leaves obtuse
at base.................................................................................. 54. L. tubulosus
Cupules enclosing the nut completely, except the umbo
Lamellae pointed, with narrowly pseudospines. Infructescence with acorns
packed close together, but not fused.......................................... 57. L. wrayi
Lamellae flattened and imbricate. Infructescence with acorns solitary, with
spaces between them
Lamellae fused on lower half, the upper half free and adaxially curved...........
.......................................................................................... 22. L. fenestratus
Lamentas fused almost to apex, only a short free lobe adaxially curved..........
.......................................................................................52. L. trachycarpus
Cupules enclosing up to 1/2 of the nut
Acorns stalked, cupules enclosing only base of the nut
Stalk up to 1 cm long. Leaves glaucous on lower surface, petiole up to 1 cm
long.................................................................................... 49. L. sundaicus
Stalk not less than 1 cm long. Leaves pale on lower surface, not glaucous,
petiole not less than 1 cm long............................................ 31. L. magneinii
Acorns sessile, cupules enclosing up to 1/2 of the nut
Acorns not less than 2 by 2.5 cm
Cupules slightly obconical. Leaves oblong, acute to caudate at apex, margin
not revolute, petiole not less than 1 cm.............................11. L. cyclophorus
Cupules saucer-shaped. Leaves obovate, obtuse at apex, margin revolute, petiole up to 0.6 cm long...........................................................44. L. revolutus
Acorns up to 1.5 by 2 cm
Nuts convex at the apex
Cupules saucer-shaped to flattened and discoid. Leaves not whorled
Lamellae usually fused throughout. Leaves up to 15 cm long....38. L. pierrei
Lithocarpus orbicarpus (Fagaceae), a new species of Stone Oak...
59
58
57
60
45
Lamellae fused at base only, apices free. Leaves not less than 18 cm...............
long................................................................................. 17. L. elephantum
Cupules cup-shaped. Leaves usually whorled at the twig tips.... 58. L. xylocarpus
Nuts flattened at the apex. Cupule cup-shaped, enclosing 1/5 to 1/2 of the
nut. Leaves with unequal sides, usually curved to one side
Leaves oblong, elliptic oblong, not less than 10 by 3.5 cm, with 14–20 pairs
of lateral nerves....................................................................... 55. L. vestitus
Acknowledgements
We are grateful to staff of the Ton Pariwat Wildlife Sanctuary for assistance in the field.
We thank R. Pooma and staff of the Forest Herbarium (BKF) for access to collections.
Finally, we would like to acknowledge S. Suddee and A. Sinbumroong for advice in
identification and the preparation of this manuscript. The first author was supported by
a Postdoctoral Research Fellowship of the Chinese Academy of Sciences (2011–2014).
References
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