Lithocarpus+elegans


 * //Lithocarpus elegans //** The Spike Oak

= What is that? = toc //Lithocarpus elegans// is one among over 300 species of the //Lithocarpus// or ‘stone oaks’ genus. In Plantae Asiaticae Rariores [9], Wallich describes the species as ­— “one of the largest as well as the commonest sort of oak in Nipal, where it attains the most gigantic size. The wood is exceedingly like the English oak in colour, and most probably equals it in other respects, but the mountaineers do not esteem it much, owing as they say to its speedy decay; a circumstance owing no doubt to their employing it in its green state.” //Lithocarpus elegans// is a regionally widespread stone oak species native to Singapore. This page seeks to inform you, the reader, of the interesting characteristics, ecological roles and taxonomy of this plant.

Many are surprised to learn that there are Oak-trees in Singapore. This is perhaps because people often think of wavy edged leaves as the distinguishing feature of Oak trees. Although this is characteristic of some English and North American species, it is the exception rather than the rule. In reality, most species have entire margins [8]. The true distinguishing feature of Oak species is the ripe fallen acorn!

Should I eat it?
There are no records of //L. elegans// acorns being consumed in Singapore, but the acorns are edible. They are rarely eaten and have reportedly been consumed in the past, in times of food shortage and in the absence of better foods. The acorns are not particularly palatable as the seeds are rich in tannins, making them bitter. The tannins can be removed by soaking the acorns in water and disposing the water bath. The fruit is usually cooked but can also be eaten raw. It may be had whole but is more often dried, ground into a powder and used for thickening stews, with cereals or to make bread [4][14].

Well, what else is it good for?
Like the fruits, the //L. elegans// bark is rich in tannins and can be used as a dye and as a preservative. The tree has yellow-brown sapwood. This wood is used in Indonesia to construct fence posts, mining props, shingles, boat building, and for making tool handles [6].

Although it is not a common sight in Singapore's parks and gardens, in other countries //L. elegans// is considered an ornamental plant and is utilised for streetscaping and landscaping purposes [4]. Indeed, there is potential to employ //L. elegans // to beautify the Singaporean "City in a Garden". It may arguably even be better suited in fact, as a native tree, than the exotic //Albizia saman, Swietenia macrophylla, Khaya senegalensis, Tabebuia rosea//, and //Xanthostemon chrysanthus// species each with over 10,000 individuals that dominate the Singaporean landscape [19].

=Where can I find it? =

** Ecology **
// Lithocarpus elegans // can be found in various habitats from mixed dipterocarp forests to lower montane forests and up to 1500 m altitude. The tree is able to thrive on a variety of soils and is tolerant of neutral to acidic pH levels [4]. As with all members of the Fagaceae family, the plant is associated with fungi by means of extensive ectotrophic mycorrhizal connections. Interestingly, it has been hypothesized that the pioneer settlement and rich development of the Oak-family in Southeast Asia, is related to the rich fungal flora in the region [2][4].

The tree acts as a refuge for various animals and its nutrient rich fruits are eaten by foraging animals such as pigs, monkeys, squirrels and rats in the forest. Long-tailed macaques (//Macaca fascicularis //) in particular have been observed enjoying the fallen fruits of the tree. The butterflies Darky Plushblue (//Flos anneilla anneilla//) and Bifid Plushblue (//Flos diardi capeta//) use the tree as a larval host [10].
 * Species interactions**

Studies have shown that //L. elegans// can accelerate community development in degraded sites such as those damaged due to extensive agriculture or practices such as shifting cultivation [24]. In Long Sungai Barang in Indonesia, //L. elegans// has been indicated as a significant agent in tree regeneration. This is due to a combination of the ability of the tree to grow in degraded sites, regenerate vegetatively, and produce a large number of seeds.
 * Community development**

Distribution
// Lithocarpus elegans // is widely distributed in East and South East Asia. It has been recorded in India, Burma, Thailand, Laos, Cambodia, Vietnam, China, Sumatra, Peninsular Malaysia, Singapore, Java, Borneo, and Sulawesi [6]. In Singapore, the tree is uncommon and critically endangered. Based on the records available at Singapore Botanic Gardens Herbarium (SING) it can be found at Mandai, Upper Pierce Reservoir, Bukit Timah Nature Reserve and in Nee Soon Swamp forest.

The regions in which //L. elegans// is known to occur are indicated on the map. You can zoom in to see the parts of Singapore where the tree has been identified.

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= When can I find it? = The tree is evergreen, flowering occurs irregularly but has been known to occur at least one a year, usually soon after the dry season. In Malaya and Singapore, flowering occurs in March and April. A distinct spermatic odour is known to emanate from the inflorescence and this plays a role in the insect pollination of the flowers. Fruiting is usually profuse and occurs within 3–6 months of flowering.

= How is it dispersed? = The fruits are large, heavy and cannot float. Dispersal cannot occur by means of water and beyond the forest margin dispersal is very slow. This is especially interesting given the vast distribution of //L. el////egans// and suggests that the present and historical distribution of the species was significantly influenced by the presence of land in past geological periods.

= How does it grow? = To germinate successfully, the nuts must be buried in the soil or covered by soil wash. The mode of germination is hypogeal, meaning the fleshy cotyledons of the nut remain underground. Under favourable conditions of high temperatures and humidity, germination may take place within 1–2 weeks after the acorns are shed. Young plants grow successfully in the shade, but older trees do well in the sun [5].

When the tree blossoms it hums with insects; attracting bees, beetles and flies for pollination. This is a characteristic of the tropical oak that distinguishes it from the temperate wind-pollinated species [7].

The female flower spike ripens into a fruit spike along which the acorns develop. Young acorns are covered by the developing cupule, which enlarges at a greater rate after pollination. As the fruit swells, the acorn presses outwards from the cup and emerges from it [7].



= How can I identify the species? = This species grows as a tree and can attain a height of up to 30 m tall. It has spirally arranged foliage and can be recognised by its distinctive acorns. The beautiful acorns of this tree usually occur in clusters and serve as its best identifying feature.

Species Description
So let's get technical! When a species is described, the author provides a set of diagnostic features which can be used to identify it. The characteristics illustrated through the photos below, as described by the species author Soepadmo, can be used to identify //L. elegans // [6].

To best understand a plant description it helps to be familiar with some technical terms. So before you dive into the diagnostic features, perhaps you would like to acquaint yourself with this short glossary of botanical terms [11]:
 * __ Abaxial __ : The side of the organ away from the axis. In the context of leaves, abaxial refers to the lower side of the leaf.
 * __ Acumen __ : Tapering point of the leaf.
 * __ Adaxial __ : The side of the organ towards the axis. In the context of leaves, adaxial refers to the upper side of the leaf.
 * __ Androgynous __ : In refernce to a stalk that bears both male and female parts.
 * __ Axis __ : The central line of symmetry of the leaf.
 * __ Cupule __ : Cup-shaped structures that covers the fruit.
 * __ Exserted __ : Protuding beyond the surrounding parts.
 * __ Fissure __ : A long, narrow opening due to seperation of plant parts.
 * __ Inflorescences __ : A flower cluster, or arrangment of flowers on the floral axis.
 * __ Lenticel __ : A pore in the stem through which gaseous exchange occurs.
 * __ Midrib __ : The middle and principal vein of the leaf.
 * __ Ovoid/ oviform/ ovoidal __ : Egg-shaped.
 * __ Rachis: __ The axis of a compound leaf or an inflorescence.
 * __ Revolute: __ Rolled downwards at the margin.
 * __ Scales __ : A flat outgrowth on any surface.
 * __ Sessile __ : Not stalked.
 * __ Globular/ Globose/ Subglobose __ : Spherical or globe shaped.
 * __ Veins __ : Vascular tissue (vessels) in a leaf.



Other //Lithocarpus// species in Singapore
It should be noted however that it can be difficult to identify this species based on vegetative material alone due to its similarity to other //Lithocarpus// species. The issue however is not particularly relevant to Singapore as //Lithocarpus elegans// is one of only thirteen species of //Lithocarpus// that occur here. Of these, only 10 are extant and one among them is a cultivated species according to the Checklist of Total Vascular Plant Flora of Singapore [8].

** Herbarium Sheet **

Specimens in a herbarium are dried, preserved, mounted on stiff sheets and labelled with relevant data. Labels usually include information such as date and place of collection, geographic co-ordinates, plant description, altitude and habitat description.

The specimens below were all determined by the //Lithocarpus// expert, E. Soepadmo. || ** Characteristic Acorn **

__Acorn__: Oak (//Lithocarpus// and //Quercus//) fruits are known as acorns. The acorns may be solitary along the rachis (i.e. fruiting stalk) or in clusters. __Cupule__: The cupule is an accessory woody structure that protects the fruit proper. It is usually cup shaped. The cupule may extend from the fruiting stem with a stalk or without one (i.e be sessile). The cupule is rarely glabrous (i.e smooth or free from hair), and is usually denticulate (i.e. finely toothed), scaly, spiny or lamellate. __Lamellae__: Refers to the distinct concentric circular layers visible on the cupule. __Fruit__: The indehiscent nuts (i.e. they do not split open on drying) range from being rounded to angular in cross section. The base of the fruit is flat. || ** Information about //Lithocarpus// species **

Of the 12 species apart from //L. elegans// known to occur in Singapore, //L. cyclophorus// and //L. gracilis// are now extinct and //L. rassa// is an exotic, cultivated species. The remaining extant native species are detailed below. || This acorn is solitary along the rachis. The cupule is saucer shaped with a short c. 5 mm long stalk, covered with dense simple hairs and has slightly distinct lamellae (5–7) with minutely denticulate edges. The cupule covers less than half the fruit and most of the acorn is exserted. It is glabrous but for the simple hairy apex. || __Conservation status__: A critically endangered, native tree.
 * **// Lithocarpus bennettii // (Bennett's Oak) ** [[image:bent.JPG width="395" height="499" align="left" caption="Specimen of L. bennettii at SING. (Photo by Srishti Arora)"]] ||< [[image:L. bennet.jpg width="230" height="238" align="center" caption="Acorn of L. bennettii. (Illustration by Srishti Arora)"]]

__Local distribution__: //Lithocarpus bennettii// thrives on well-drained soils of primary heath and swamp forests. It is found locally, in the Nee Soon Swamp forest.

__Interesting Note__: Though it is presently not cultivated in Singapore, the tree has potential uses in landscaping of parks and gardens. Its timber can be used. || This acorn is solitary along the rachis. The cupule is cup shaped with a c. 5–7 mm long stalk, densely hairy and has slightly distinct lamellae (5–9) with entire or denticulate edges.
 * ** //Lithocarpus cantleyanus// (Cantley's Oak) ** [[image:cantly.JPG width="396" height="503" align="left" caption="Specimen of L. cantleyanus at SING. (Photo by Srishti Arora)"]] || [[image:L. cant.jpg width="207" height="345" align="center" caption="Acorn of L. cantleyanus. (Illustration by Srishti Arora)"]]

The cupule covers less than half the fruit and most of the acorn is exserted. It is glabrous. The fruit base is rounded and the apex is abruptly acute. || __Conservation status__: A critically endangered, native tree.

__Local distribution__: //Lithocarpus cantleyanus// thrives on clay soils of primary forests. The tree is cultivated in Singapore.

__Interesting Note__: The species is named for a superintendent of the Singapore Botanic Gardens, Nathaniel Cantley. || This acorn is solitary along the rachis and distantly spaced. The cupule is cup shaped with a c. 3–8 mm long stalk, densely hairy and has slightly distinct lamellae (5–6) with denticulate edges.
 * ** //Lithocarpus conocarpus// (Singapore Oak) ** [[image:conocarpu.jpeg width="397" height="680" align="left" caption="Specimen of L. conocarpus at L. (Photo from Naturalis Bioportal)"]] || [[image:L. cono.jpg width="186" height="287" align="center" caption="Acorn of L. conocarpus. (Illustration by Srishti Arora)"]]

The cupule covers up to half the fruit and most of the acorn is exserted. The fruit is hairy. The fruit base is rounded and the apex is abruptly acute. || __Conservation status__: A critically endangered, native tree.

__Local distribution__: //Lithocarpus conocarpus// thrives on clay-rich soils of primary forests. The tree is cultivated in Singapore.

__Interesting Note__: The species was once quite common in Singapore, hence its common name — Singapore Oak. || This acorn is solitary along the rachis but arranged closely together. The cupule is deeply cup shaped, covering almost the entire fruit with a c. 10–15 mm long stalk, densely hairy and has slightly distinct lamellae (5–7) with minutely denticulate or entire edges.
 * **//Lithocarpus encleisacarpus// (Covered Oak)** [[image:encleisacarpus.jpg width="392" height="509" align="left" caption="Specimen of L. encleisocarpus at SING. (Photo by Srishti Arora)"]] || [[image:L. encleisa.jpg width="226" height="300" align="center" caption="Acorn of L. encleisacarpus. (Illustration by Srishti Arora)"]]

The cupule covers most of the acorn, the exserted fruit is velvety. || __Conservation status__: A critically endangered, native tree.

__Local distribution__: //Lithocarpus encleisacarpus// is found in primary and lower montane forests.

__Interesting Note__: The species can be recognized from its silky acorns covered by the thin cup with distinct rings. || This acorn is solitary along the rachis. The cupule is cup shaped, covering less than half the fruit with a c. 5–10 mm long stalk, sparsely hairy and has slightly distinct lamellae (6–7) with entire edges.
 * ** //Lithocarpus ewyckii// (Ewijk's Oak) ** [[image:ewyckii.jpg width="392" height="501" align="left" caption="Specimen of L. ewyckii at SING. (Photo by Srishti Arora)"]] || [[image:L. ewyickii.jpg width="194" height="264" align="center" caption="Acorn of L. ewyckii. (Illustration by Srishti Arora)"]]

Most of the ovoid fruit is exserted with a rounded base and abruptly acute apex. || __Conservation status__: An endangered, native tree.

__Local distribution__: //Lithocarpus ewyckii// found on sandy loam and clay soils in primary and lower montane forests. The tree is cultivated in Singapore. || This acorn is sessile and solitary along the rachis. The cupule is saucer shaped, covering less than half the fruit, irregularly set with slender prickle like appendages.
 * ** //Lithocarpus hystrix// (Porcupine Oak) ** [[image:hystrix.jpeg width="391" height="526" align="left" caption="Specimen of L. hystrix at SING. (Photo by Srishti Arora)"]] || [[image:L. hystrix.jpg width="238" height="215" align="center" caption="Acorn of L. hystrix. (Illustration by Srishti Arora)"]]

Most of the ovoid or conical glabrous fruit is exserted with a rounded base and abruptly acute apex. || __Conservation status__: A critically endangered, native tree.

__Local distribution__: //Lithocarpus hystrix// found on sandy-clay soils in primary and lower montane forests. The tree is cultivated in Singapore.

__Interesting Note__: The species is named for the slender prickles that cover its cupule. || This acorn is sessile or with a stalk c. 5 mm and solitary or in clusters of three along the rachis. The cupule is cup to saucer shaped, covering less than half the fruit, the lamellae are dense and distinct set in 8–11 lines.
 * ** //Lithocarpus lucidus// (Shining Oak) ** [[image:lucidus.jpeg width="387" height="668" align="left" caption="Specimen of L. lucidus at L. (Photo from Naturalis Bioportal)"]] || [[image:L. lucid.jpg width="262" height="216" align="center" caption="Acorn of L. lucidus. (Illustration by Srishti Arora)"]]

Most of the depressed ovoid glabrous shining fruit is exserted with a rounded base and rounded apex. || __Conservation status__: An endangered, native tree.

__Local distribution__: //Lithocarpus lucidus// found on well drained soils in primary and lower montane forests. It occurs locally in Nee Soon Swamp Forest and the Upper Seletar Reservoir and Upper Pierce Reservoir areas. The tree is cultivated in Singapore.

__Interesting Note__: The species is named for its shiny fruit. || This acorn is sessile or with a stalk c. 5–10 mm and solitary along the rachis. The cupule is saucer shaped, densely hairy, with distinct scaly appendages set in regular lines.
 * ** //Lithocarpus sundacius// (Sunda Oak) ** [[image:sundac.jpeg width="384" height="658" align="left" caption="Specimen of L. sundacius at L. (Photo from Naturalis Bioportal)"]] || [[image:L. sunda.jpg width="216" height="262" align="center" caption="Acorn of L. sundacius. (Illustration by Srishti Arora)"]]

Most of the depressed ovoid glabrous shining fruit is exserted and has a hairy apex. || __Conservation status__: A critically endangered, native tree.

__Local distribution__: //Lithocarpus sundacius// is found on well drained soils in primary and lower montane forests. The tree is cultivated in Singapore.

__Interesting Note__: The species is named for its occurrence on the Sunda Islands. || This acorn is sessile and in closely arranged clusters of 2–3 along the rachis. The cupule is saucer shaped, densely hairy, with distinct scaly appendages set in irregular lines.
 * ** //Lithocarpus wallichianus// (Wallich's Oak) ** [[image:wall.jpeg width="381" height="662" align="left" caption="Specimen of L. wallichianus at L. (Photo from Naturalis Bioportal)"]] || [[image:L. wallich.jpg width="245" height="245" align="center" caption="Acorn of L. wallichianus. (Illustration by Srishti Arora)"]]

Most of the ovoid fruit is exserted and is covered with simple hairs. || __Conservation status__: A critically endangered, native tree.

__Local distribution__: //Lithocarpus wallichianus// is found on well drained soils in hilly forests. The tree is cultivated in Singapore.

__Interesting Note__: The species closely resembles the Common Oak (//L. sundacius//) especially the fruits, but has hairy twigs, larger leaves and longer fruit spikes with many acorns on them. ||

 media type=custom key=2944217 = What is it called again? =

Taxonomy
Kingdom: Plantae Order: Fagales Family: Fagaceae Genus: //Lithocarpus// Species: //Lithocarpus elegans//

Fagaceae
Though the Fagaceae dominate in temperate forests, the center of diversity is in Southeast Asia. //Lithocarpus// in particular are restricted to Asia and no species are known to exist in the Northern hemisphere.

By Soepadmo’s admission the classification of Fagaceae species is based largely on gross morphological attributes, in particular the taxonomic limits are based on fruit and floral characteristics such as pollination syndrome and number of cupule valves.. Some systematists have proposed splitting the family into two distinct ones, or to split a few genera such as //Quercus// and //Trigonobalanus// into smaller genera. Even at the subfamily level, proposals have been made to rearrange the genera into different subfamilies and tribes. These changes are not yet widely accepted and for the most part the classification prescribed by Kubitzki (1993) is still used [6].

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 13px;">The family Fagaceae currently includes nine genera— //Fagus, Castanea, Castanopsis, Chrysolepis, Colombobalanus, Formanodendron, Lithocarpus, Quercus,// and //Trigonobalanus// [25].

//Lithocarpus//
In the past, there has been disagreement among botanists regarding whether the genera //Lithocarpus// and //Quercus// are sufficiently different and on whether //Lithocarpus// should be subdivided into at least 2 or 3 smaller genera [5].

Lithocarpus was divided by Oersted (1867) into //Cyclobalanus// and //Pasania.// Similarly, Schwarz (1936) preferred to split the genus into three, //Lithocarpus//, //Cyclobalanus,// and //Pasania.// Oersted and Schwarz cited distinct cupule characteristics as a means of distinction for the smaller genera they recognized. Soepadmo indicates that generic distinction on the basis of one character is “always unsatisfactory”. Similarly in the case of //Lithocarpus// he argues that a narrow generic limit is invalid because there are many species in which cupule morphology is unclear. He also raises issue with the fact that there is general “uniformity in floral structure, pollen morphology, wood anatomy, and floral biology” for the //Lithocarpus// genus. Any one character is therefore not sufficient for distinction smaller genera. Soepadmo’s recommendation is that the “narrow generic concept be rejected” [5].

Soepadmo's position is in line with some past authors. Prantl (1889), placed all species in a single genus //Pasania//. Koidzumi (1916), likewise agreed to include all species in one genus, but under the name //Synaedrys//. Rehder & Wilson in 1916 restored //Lithocarpus// as the correct name for the genus [5].

Nomenclature
**Scientific name:** //Lithocarpus elegans// The genus name '//Lithocarpus'// is derived from the Greek '//lithos'// for stone and '//carpos'// meaning fruit, in reference to the hard acorns. The specific epithet '//elegans'// is Latin for elegant, referring to the elegant acorns and cupule. **Author citation**: // Lithocarpus elegans // (Blume) Hatus // ex. // Soepadmo This author citation is complex and reflects how the species has been classified over the years. In the parenthesis is the name of the first author, C. L. Blume, who had assigned the specific epithet in a different genus (//Quercus//). It was then moved to the present genus (// Lithocarpus) //by the revising author S. Hatusima. The //'ex.'// in the citation denotes that the subsequent author E. Soepadmo provided the validating description for the species. **Vernacular names:** The tree is known by different vernacular names in different regions. In Singapore it is also known as Spike oak, Mempening, Empening or Berangan.

Protologue
The first valid published material associated with a scientific name, including diagnosis, description and illustrations constitutes the protologue. On an unrelated but interesting note, in the protologue material on this species the long 's' ( ſ) is used in the publication. This is similar in appearance to the minuscule 'f'. Which is why to the modern reader the description may seem //' ſ trange'//.

This plant was first described as //Quercus spicata// by Smith in the Cyclopaedia (1809). This was the first of many names attributed to the this widely distributed and highly variable tree (See Synonyms). The species has been moved back and forth across genera. In the past, over seven varieties of the tree have been distinguished and some have even been recognised as distinct species. It was only in 1970 that Soepadmo concluded that none of the varieties deserved to be regarded as a separate species. He described //Lithocarpus elegans// as the most primitive of the genus and argued that the variations in morphology of the plant are primarily due to growth in different habitats and that in the past poor characters had been used to delimit the species. He instead suggested that the //Lithocarpus elegans// species be recognised by the "glabrous, pale to dark chocolate brown leaves, glossy on both sides and by the cupules which are always squamous and set in dichasial clusters of 3–10" [4]. Based on his treatment of the species //L. elegans// and the genus //Lithocarpus// it could be argued that Soepadmo ascribes to a relatively broad species concept. The dramatic variation in the species is apparent in the images below, all of which are specimens from Peninsular Malaysia (accessed at SING) and have been determined as //Lithocarpus elegans// by species author Soepadmo.

**Basionym**
The term basionym refers to the original name on which the present name attributed to the species is based. For //Lithocarpus elegans,// the basionym is //Quercus elegans// Blume.

A rough translation of the latin description — The leaves are ovate-oblong, bluntly pointed with entire margin, smooth petiole, dense fruit with a hemispherical nut.

Synonyms
<span style="font-family: Arial,Helvetica,sans-serif;">Synonyms are names that have been rejected by a particular author or authors. A synonym may be rejected for either of two reasons: (1) because it does not comply with the rules of the International Code of Botanical Nomenclature (ICBN); or (2) because a particular author rejects the classification associated with the synonym [23]. This is a selection of synonyms of //L. elegans// in Malesia. However there are even more synonyms, the full list can be found in Soepadmo's Florae Malesianae Praecursores XLIX (1970).

// Q. spicata // (//non// Humb. & Bonpl. 1809) Smith //in// Rees, Cyclop. (1814) 12; //Q. racemosa// (//non// Lam. 1783) Jack, Mal. Misc. 2 (1822) 86; //Q. arcaula// Hamilt. //ex// G. Don var. //racemosa// (Jack) Blume //l.c.// (1850) 290; //Q. spicata// (//non// Humb. & Bonpl.) Smith var. //racemosa// (Jack) Miq., Ann. Mus. Lugd.-Bat 1 (1863) 106; //Q. depressa// Blume //l.c.// (1823) 209, //t.// 1; //Q. spicata// (//non// Humb. & Bonpl.) Smith var//. depressa// (Blume) King, Ann. Roy. Bot. Gard. Calc. 2 (1889) 48, //t.// 43; //Q. glaberrima// Blume //l.c.// (1823) 210, //t.// 2; //Q. spicata// (//non// Humb. & Bonpl.) Smith var//. glaberrima// (Blume) Miq. //l.c.// (1856) 848; //L. spicata// (Smith) Rehder var//. glaberrima// (Blume) A. Camus //l.c//. (1949) 103, //t.// 482; //L. spicata// (Smith) Rehder var. //elegans// (Blume) A. Camus //l.c.// (1949) 102, //t.// 481; //Q. placentaria// Blume, //l.c.// (1825) 518; //Q. spicata// (//non// Humb. & Bonpl.) Smith var. //placentaria// (Blume) Schottky //l.c.// (1912) 664; //Q. grandifolius// D. Don //in// Lamb., Gen. Pin. 2 (1824) 27, //t.// 8; //L. grandifolius// (D. Don) S.N. Biswas, Bull. Bot. Surv. India 10 (1968) 258; //P. spicata// (Smith) Oerst. var. //placentaria// (Blume) Oerst. //l.c.// (1867) 83; //Pasania spicata// (Smith) Oerst. var//. placentaria// (Blume) Schottky //l.c.// (1912) 664; //L. spicata// (Smith) Rehder var//. placentaria// (Blume) Rehder //l.c//. (1929) 133; //Q. glomerata// Roxb., Fl. Ind. 3 (1832) 640; //P. glomerata// (Roxb.) Oerst. //l.c.// (1891) 379; //Q. anceps// Korth. //l.c.// (1844) 204; //Q. microcalyx// Korth. //l.c.// (1844) 206; //Q. arcaula// Hamilt. //ex// G. Don var//. microcalyx// (Korth.) Blume //l.c.// (1850) 290; //Q. spicata// (//non// Humb. & Bonpl.) Smith var. //microcalyx// (Korth.) Miq. //l.c.// (1856) 848; //P. spicata// (Smith) Oerst. var. //microcalyx// (Korth.) Gamble //l.c.// (1915) 421; //L. microcalyx// (Korth.) A. Camus //l.c//. (1945) 83; //Q. pyrifolia// Blume //l.c.// (1850) 304; //Q. sphacelata// Blume //l.c//. (1850) 304; //Q. gracilipes// Miq., Fl. Ned. Ind. 4, Suppl. (1861) 347; //Q. spicata// (//non// Humb. & Bonpl.) Smith var//. gracilipes// (Miq.) Miq. //l.c.// (1863) 106; //P. spicata// (Smith) Oerst. var//. gracilipes// (Miq.) Schottky //l.c.// (1912) 664; //L. spicata// (Smith) Rehder var//. gracilipes// (Korth.) Rehder //l.c.// (1919) 131; //Q. spicata// (//non// Humb. & Bonpl.) Smith var//. latifolia// Scheff. //l.c.// (1870) 359; //Q. rhioensis// Hance //l.c.// 198; //L. rhioensis// (Hance ) A. Camus //l.c.// (1932) 40.

Type specimen
The International Code of Botanical Nomenclature (ICBN) states that scientific names need to be associated with a physical entity known as a nomenclatural type or simply type. The nomenclatural type is usually a herbarium sheet specimen; other specimens of the species are compared with the type specimens [20].

The __holotype__ is the specimen designated in the original publication of the species on which the name is based. The Holotype for the species collected by Blume from Bantam Residency in Java is housed at the Leiden branch of the National Herbarium of the Netherlands (L). An __isotype__ is a duplicate specimen of the holotype collected at the same time and from the same population. It is recommended that isotypes be designated in the publication of a new name as they are reliable duplicates. Isotypes of //L. elegans// are available at the Herbarium at Royal Botanic Gardens Kew (K), the herbarium of Muséum National d'Histoire Naturelle (P) and at the Cambridge University Herbarium (CGE).
 * Types of types**

Other types of types include [20]: __Lectotype__: A specimen from the original publication that shall serve as the type if no holotype is designated at the time of publication. __Neotype__: A specimen that is not included in the original material that shall serve as the type if all original material is missing. __Syntype__: Multiple specimens all designated as types, or a specimen cited in the absence of a holotype. __Paratype__: All specimens not designated as holotypes, isotype or syntype but cited in the original material. Soepadmo has cited several specimens in his publication for // L. elegans. // __Epitype__: A specimen designated to serve as the type if the holotype, lectotype or neotype is ambiguous,



Phylogeny
There have not been many phylogenetic studies of the genus //Lithocarpus//, and no published accounts could be found for //L. elegans//. The most comprehensive analyses available are the ‘Systematics of Fagaceae‘ by Manos et al (in which the authors used DNA sequence data to reconstruct phylogeny, assess systematic relationships, and explore alternative patterns of morphological specialization) and ‘Morphological and molecular diversity in Lithocarpus of Mount Kinabalu’ by Cannon et al [25] [26].

For the most part, taxonomic classification of //Lithocarpus// has been stable, forming the most strongly supported group (bootstrap analysis) in the Fagaceae family and the genus appears to be monophyletic. In the genus, the species are divided into distinct sections based on fruit morphology. Manos et al’s results indicate that several sub-clades in //Lithocarpus// corresponded to the previously delimited groups [26][27].

It is interesting to note that in general the species of //Lithocarpus// tend not be widespread making //L. elegans// and //L. grandifolius// notable exceptions. Unfortunately, this aspect of //L. elegans// phylogeography specifically has not been explored. It has been indicated that initially the ancestral group of //Lithocarpus// was widespread across South and Southeast Asia allowing the ancestral haplotype to reach either end of the subcontinent without significant alteration. Subsequently, the widespread population became fragmented and experienced an extreme bottleneck in central Sundaland. This bottleneck together with fragmentation caused the Bornean lineage to undergo significant genetic drift and to diversify in isolation, which may explain why the greatest diversity of haplotypes is found in Borneo and the relatively low frequency of occurrence of the ancestral haplotype. It is safe to say however that species boundaries of //Lithocarpus// are yet to be fully understood and further phylogenetic analyses are likely to reveal this [25] [26].

All photos are used with permission from photographers unless otherwise stated. Credit is attributed to photographers in the picture caption. For the composite image at the top of the page: The photos were taken by Srishti Arora, Louise Neo and Reuben Lim (From left to right).

=**References**=

(Accessed from the Biodiversity Heritage Library on 1 November 2017) ||
 * [1] Cannon CH & Manos PS (2001) The Bornean Lithocarpus Bl. section Synaedrys (Lindl.) Barnett (Fagaceae): its circumscription and description of a new species. Botanical Journal of the Linnean Society, 133: 343–357. ||
 * [2] Corner EJH (1988) Wayside Trees of Malaya. The Malayan Nature Society, 1: 327–343. ||
 * [3] Julia S & Soepadmo E (1998) New Species and New record of Lithocarpus Blume (Fagaceae) from Sabah and Sarawak, Malaysia. Gardens' Bulletin Singapore, 50: 125–150. ||
 * [4] Soepadmo E (1972) Flora Malesiana. Wolters-Noordhoff Publishing, Groningen, The Netherlands, pp. 265–403. ||
 * [5] Soepadmo E (1970) Florae Malesianae Praecursores XLIX Malesian Species of Lithocarpus Bl. (Fagaceae). Reinwardtia, 8(1): 197–308. ||
 * [6] Soepadmo E, Julia S & Go R (1995) Fagaceae. In: Soepadmo E & Wong KM (ed.) Tree Flora of Sabah and Sarawak. Volume 1. Forest Research Institute of Malaysia (FRIM), Sabah Forestry Department, and Sarawak Forestry Department, Kuala Lumpur, Malaysia, pp. 1–166. ||
 * [7] Whitmore TC (1972) Tree Flora of Malaya A Manual for Foresters. Longman, Forest Research Institute, Kepong, 1: 198–209. ||
 * [8] Chong KY, Tan HTW & Corlett RT (2009) A CHECKLIST OF THE TOTAL VASCULAR PLANT FLORA OF SINGAPORE: Native, Naturalised and Cultivated Species. Raffles Museum of Biodiversity Research, National University of Singapore, pp. 56. ||
 * [9] Treuttel and Wurtz (1832) Plantae Asiaticae rariores, or, Descriptions and figures of a select number of unpublished East Indian Plants. London, pp. 40–41.
 * [10] Tan H (2012) Butterflies of Singapore. Version 1 December 2017. [] (Accessed 1 December 2017) ||
 * [11] Hickey M and King C (2000) The Cambridge Illustrated Glossary of Botanical Terms. Cambridge University Press, pp. 1–47 ||
 * [12] Colin Mills (2010) Lithocarpus elegans (Blume) Hatus ex. Soepadmo. Version 28 February 2010. [] (Accessed 1 December 2017) ||
 * [13] National Parks Board (2013) Flora and Fauna Web:Lithocarpus elegans (Blume) Hatus ex. Soepadmo. [] (Accessed 1 December 2017) ||
 * [14] Ken Fern (2014) Useful Tropical Plants Database: Lithocarpus elegans. [] (Accessed 1 December 2017) ||
 * [15] Longman, Hurst, Rees, Orme & Brown (1819) The cyclopaepedia; or, Universal dictionary of arts, sciences and literature. University of California Libraries, 29. (Accessed from the Biodiversity Heritage Library on 1 November 2017) ||
 * [16] Biodiversity Heritage Library (2017) Biodiversity Heritage Library. [] (Accessed 1 December 2017) ||
 * [17] Bioportal (2017) Naturalis Biodiversity Centre. [] (Accessed 1 December 2017) ||
 * [18] JSTOR (2017) Global Plants Database. [] (Accessed 1 December 2017) ||
 * [19] Nghiem LTP, Tan HTW and Corlett RT (2015) Invasive trees in Singapore: are they a threat to native forests? Tropical Conservation Science, 8(1): 201–214. ||
 * [20] Simpson MG (2006) Plant Nomenclature: Plant Systematics. Elsevier Academic Press, pp. 501–517. ||
 * [21] Simpson MG (2006) Plant Nomenclature: Phylogenetic Systematics. Elsevier Academic Press, pp. 17–51. ||
 * [22] Simpson MG (2006) Plant Nomenclature: Diversity and Classification of Flowering Plants: Eudicots. Elsevier Academic Press, pp. 227–347. ||
 * [23] Simpson MG (2006) Plant Nomenclature: Resources in Plant Systematics. Elsevier Academic Press, pp. 507. ||
 * [24] Cairn MF (2015) <span style="background-color: #ffffff; font-family: Arial,sans-serif;">Shifting Cultivation and Environmental Change: Indigenous People, Agriculture and Forest Conservation. Routledge, pp.425–428. ||
 * [25] Manos PS et al (2001) Systematics of Fagaceae - Phylogenetic tests of reproductive trait evolution. International Journal of Plant sciences, 162(6): 1361–1379. ||
 * [26] Cannon HC and Manos PS (2003) Phylogeography of the southeast asian stone oaks Lithocarpus. Journal of Biogeography, 30: 211–226. ||
 * [27] Cannon HC (2001) Morphological and molecular diversity in Lithocarpus of Mount Kinabalu. Sabah Parks Nature Journal, 4: 45–69. ||