Fagaceae

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Fagaceae

Description

Monoecious trees or rarely shrubs, in Mal. evergreen, sometimes buttressed or with stilt-roots; — In Malesia most species of Fagaceae are trees of medium to large size, rarely shrubs of c. 2-5 m tall (Lithocarpus orbicularis, L. oreophilus, Nothofagus pullei). Buttresses and stilt-roots are often present in several species of Lithocarpus and Quercus.
— Most Malesian species of Fagaceae are readily recognizable in the field by their characteristic light-greyish, smooth and mottled bark; only a few have bark which is grey-brown and rather rough. In Nothofagus the bark is often peeling in large flakes. Lithocarpus, Nothofagus, Quercus and Trigonobalanus verticillata are very easily recognized by the same typical slash characters: a reddish-brown bark, easily detached from the smooth whitish wood, with numerous longitudinal faintly bluish lines about ½-1½ cm long and 1 mm wide and slightly depressed. The opposite parts on the inside of the bark are correspondingly slightly elevated. This typical pattern is not found in Castanopsis.
Hairs simple or stellate or fasciculate, rarely with resiniferous colleters, or scales on pits on the underside of the leaf. — The leaves of Malesian Fagaceae are only very rarely not entire; they are at most crenate or sinuate near the upper half in some species of Castanopsis and Quercus, and remotely, shallowy crenate in a few species of Nothofagus. Domatia are not found in Malesian Fagaceae.
Leaves simple, spirally arranged, rarely in whorls of 3 or distichous, sometimes crowded near the top of each flush, penninerved, in Mal. entire or rarely crenate or sinuate. Stipules present, caducous or rarely rather long persistent, rarely interpetiolar or peltately attached. — Castanopsis, Lithocarpus, and Quercus have extra-petiolar, caducous stipules, and Nothofagus possesses stipules which are distinctly peltately attached and in the flush they develop earlier than the leaves. The stipules in Trigonobalanus verticillata are extra-petiolar in the first few leaves of the seedling, and later become interpetiolar.
Inflorescence a cyme or a simple or branched spike, bracteate, ♂, ♀, androgynous (with the ♀ flowers borne on the lower part) or mixed. — A pistillode is present in the ♂ flower of Castanea, Castanopsis, and Lithocarpus, and replaced by a cluster of simple stiff hairs in Trigonobalanus verticillata and some species of Malesian Quercus. In the ♀ flowers 6-12 staminodes are found in the above-mentioned genera. In some species of Lithocarpus (L. ruminatus and L. turbinatus) from Borneo, they are rather well developed and exceed in length the height of the perianth-lobes. The stigma is capitate in Malesian Quercus and Trigonobalanus, punctiform in Castanea, Castanopsis, and Lithocarpus, or forming a broad stigmatic surface on the inner side of the style-arms (Fagus, Nothofagus and extra-Malesian Quercus).
Flowers unisexual or functionally so. Fruit an indehiscent nut (achene), 1-3-celled, sometimes falsely multiseptate, rounded or sharply 2-3-an-gular. Seed one, exalbuminous;

Distribution

77°S: Africa: present Am Is: absent America: present Antarctic:, Antarctica (Antarctica) Asi: present Asia-Temperate:, Manchuriaabsent Asia-Tropical: Australasia:, Tasmania (Tasmaniapresent) Bohemia: Ceylon: absent Deccan Peninsula: absent Europe:, Iceland (Iceland) Greenland: In the Old World the distribution extends southwards from 62°N in Scandinavia southheastwards to Kashmir and then northeastwards to the Sea of Okhotsk at c. 55°N: present Japan: Kangean Is: absent Key Is: absent Lesser Sunda Is: absent Madura: absent New Caledonia: present New Ireland: absent New Zealand: North America: Northern America:, Colorado (Colorado); Montana (Montana); Wyoming (Wyoming) SE. Australia: SE. Deccan Peninsula: Seymour I: Silesia: Solomons: absent South America they occur from Fuegia and Staten I. northwards to Argentina and on the western slopes of the Andes in Chile up to 33°S: present Tanimbar Is: absent Vancouver I: W. Germany: Woodlark I: absent almost the whole of Australia: dry parts of the Middle East: absent extreme northern parts of Japan: absent from Canada and the United States southwards to Central America, as far south as a few scattered localities in Columbia, in South America: present from the Malay Peninsula eastwards to d'Entrecasteaux Is., New Britain, and the Louisiades to c. 153°E: present northern hemisphere: northern rim in the western Mediterranean region: present otherwise absent from Pacific islands: absent scarce wet parts of East Australia: present seasonally dry parts of Central and East Java: present southern Central Asia: southern Moluccas: absent southern hemisphere: present the desert and colder parts of China: absent western part of North America extending to 65°N:

Dispersal

— Since the fruits are not provided with any special means of dissemination and are generally fairly large, mostly rounded, and fairly heavy, dispersal in this family must be slow. The fruits are not capable to float either in freshwater or in the sea. Not far from river-mouths the tropical beach is often strewn with acorns, but these are barren, hence buoyant and drifted downstream to the ocean (cf. ). This is in complete accordance with the experiments and observations made by PREEST on Nothofagus in New Zealand (); he found that the fruits do not travel very far from the stands and are poorly fitted for dispersal beyond the forest margin. A similar observation was made by KALKMAN & VINK in New Guinean Nothofagus (). This slow dispersal makes the distribution of the family most interesting, suggesting that their present and fossil distribution is an indicator of prime importance for the presence of land in former geological epochs. See also .

Taxonomy

Historical review. — DUMORTIER () was the first to recognize the family Fagaceae, and A. DE CANDOLLE () was the first to frame the family in its present circumscription, using the name Cupuliferae. OERSTED () accepted DE CANDOLLE'S classification and used characters of the stigma, cotyledons, and germination to subdivide the family into three subfamilies: Quercineae to include Quercus L. and Cyclobalanopsis OERST.; Castanineae: Pasa-nia (MIQ.) OERST., Cyclobalanus (ENDL.) OERST. (including Lithocarpus Bl.), and Castanea MILL. (including Castanopsis SPACH); and Fagineae: Fagus L. and Nothofagus Bl.

PRANTL (), who first used the name Fagaceae and recognized the importance of the inflorescence, divided the family into two tribes, viz Fageae: Nothofagus and Fagus; and Castaneae: Castanea (including Castanopsis),Pasania (includingLithocarpus and Cyclobalanus), and Quercus (including Cyclobalanopsis).

O. SCHWARZ (), who combined the characters already mentioned by OERSTED and PRANTL, and added those of the cupule and fruit, came to the same subdivision as OERSTED. However, he recognized more genera than his predecessors, namely by splitting the genus Quercus into four segregates: Quercus, Cyclobalanopsis, Erythroba-lanus O. SCHWARZ, and Macrobalanus O. SCHWARZ; and Lithocarpus into three: Cyclobalanus, Lithocarpus, and Pasania.

Among the recent authors who basically accept OERSTED'S classification, though they used different combinations of characters, are: MELCHIOR () and FORMAN (); those who agree with PRANTL are: BRETT () and LUONG ().

On the basis of her pollen-morphological study, KUPRIANOVA () recently suggested to accommodate the genera Nothofagus and Trisyngyne BAILL. into a distinct family, Nothofagaceae, and place this new family in the Euphorbiales. Her opinion is, however, completely unacceptable to us, as we found that the resemblance between the pollen of Nothofagus (including Trisyngyne) and that of Longetia (Euphorbiaceae) is highly superficial. Furthermore, the presence of a cupule containing fruits which are nuts, is the most important feature characterizing the family Fagaceae, and is nowhere found among the Euphorbiales. Moreover, the morphology of Nothofagus fits admirably in with that of Fagaceae especially Fagus and not at all in with that of Euphorbiaceae. It is true that the pollen of Nothofagus is morphologically very different from that of the rest of the family. Morphologically it is even so isolated, that Mr. J. MULLER admits that if only the pollen was known, a palynologist would be at a loss to place it in any family! However, the problem in Nothofagus stands not alone, as there are many genera or families in which the pollen is very highly variable morphologically, and yet they are accepted as belonging to a single larger taxon.

Cytology

Data on tropical and subtropical species are unfortunately scant. In various extra-Malesian species diploid numbers have been reported to be 20 or 24 (Castanea, Fagus, Quercus) and recently 28 in Quercus castaneifolia CAMUS (TUTAJUK et ah,). ARMSTRONG & WYLIE reported for Nothofagus 2n = 26. Independently KWITONG JONG (in litt.) and DING HOU found for Trigonobalanus verticil lata 2n = 44 (DING HOU also n = 22). As to chromosome number Fagaceae have a great affinity with that found in other Amentiferous orders (Betulaceae, Corylaceae).

Uses

In comparison with the importance of oakwood, oak tannin, beechwood, and chestnuts in the northern temperate regions, the economic value of Fagaceae in Malesia is slight, Nothofagus obviously excepted. The wood of Castanopsis, Lithocarpus and Quercus is often very hard and difficult to work, although it is sometimes rather beautifully and evenly grained. It is very liable to splitting, and seems rather unfit for construction purposes, for which it is occasionally used in Indonesia. Records of durability vary with the species. Some New Guinean species are supposed to be promising for furniture and veneer, but tests have not been made, and species may differ considerably in properties.

According to field notes several Nothofagus species, notably N. starkenborghii, N. perryi, N. pullei, and N. rubra, possess excellent very resistant hardwood, used for bridge constructions and locally used on a large scale also for general constructions under roof. The Papuans in East New Guinea plant various species (N. grandis, N. pullei, N. perryi) in their native homesteads, around their villages, along tracks and around their garden lands, it is said for ornamental purpose, but probably also intended for later use. For this purpose they collect seedlings in the forest.

In Borneo, Dr. J. A. R. ANDERSON informed me, that logs of some species of Castanopsis, Lithocarpus, Quercus, and Trigonobalanus verticillata are recently being tried for mushroom cultivation. This trial was apparently inspired by the success of mushroom cultivation on wood of Quercus species in Japan and made possible by the occurrence of closely allied edible mushrooms on Mt Kinabalu.

The fruits of some species of Castanopsis (e.g. C. argentea, C. costata, C. inermis, C. javanica, and C. tungurrut) are widely consumed after cooking or roasting, just like chestnuts in Europe and other temperate regions.

The bark in many species of Fagaceae contains appreciable amounts of tannin, but perhaps because of the hardness of the wood, there seems to be no commercial exploitation of it. See HEYNE () and BURKILL () for details.

Phytochemo

All Fagaceae seem to be accumulators of polyphenolic compounds; flavonols, leucoanthocyanins, catechins and gallic and ellagic acid occur frequently in leaves, fruits, bark and wood. In wood of Nothofagus species flavanones (naringenin), flavononols (aromadendrin, taxifolin), stilbenes (pinosylvin, resveratrol) and dihydrochalcones (nothofagin, konnanin) have been observed in addition (). Besides these low molecular polyphenolic compounds most members of the family contain rather large amounts of tannins. Oak bark, oak wood, oak gall-nuts and chestnut wood represent important sources of vegetable tannins. Oak trees elaborate predominantly galli- and ellagitannins in leaves, gall-nuts and wood and predominantly condensed tannins in barks. The same may be true for other members of the family. Hamamelitannin, a simple gallitannin. accompanies condensed tannins in the bark of Castanea sativa and Quercus rubra. Turkish tannin is the gallitannin of Aleppo gall-nuts. Castalgin, vescalgin, castalin and vescalin are complex ellagitannins of the wood of Castanea sativa and Quercus petraea. Pedunculagin is an ellagitannin of known structure from gall-nuts of Cental European species of Quercus. Besides polyphenolic compounds members of Fagaceae contain rather large amounts of pentacyclic triterpenes in barks, leaves and gall-nuts. Friedelin and the two epimeric friedelanols have been observed most frequently but derivatives of α- and β-amyrin as well as still other triterpenes do occur also. All species of the large genus Quercus seem to store quercit in leaves, bark and seeds; this represents a character of the genus Quercus. In conclusion it may be stated that Fagaceae resemble other Amentiferous families in tannin-accumulation, in the patterns of low-molecular phenolic compounds and in the tendency to produce conspicuous amounts of triterpenes. Chemistry also favours a relationship with the Hamamelidaceous and Rosaceous stock.
General reference: . — R. HEGNAUER.

Embryology

So far no study has been made on the Malesian species of Fagaceae. HJELMQVIST basing his observations on Quercus robur, Fagus sylvatica, and Castanea sativa concluded that the development of the endosperm and the embryo in Fagaceae is rather similar to that of Juglandaceae, but differs very sharply from that of Betulaceae.