Chloranthaceae

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Chloranthaceae

Description

Erect or straggling herbs, shrubs or trees, sometimes monoecious or dioecious, the herbs sometimes rhizomatous; Leaves simple, decussate or sometimes whorled in fours, serrate, crenate or dentate, the teeth often thickened at the apex, penninerved, usually petiolate; Flowers much reduced, without perianth, fully unisexual or essentially bisexual with the reduced anther-bearing organ adnate to the side of the ovary; Ovule solitary, orthotropous, pendulous, bitegmic and crassinucellate. Seeds subglobose, exarillate, with copious fleshy or oily endosperm and minute embryo, the cotyledons divaricate or scarcely formed.

Distribution

Africa: present Asia-Temperate:, Koreapresent; Palestine (Israelpresent) Asia-Tropical:, Borneopresent; Indiapresent; New Guineapresent; Philippines (Philippinespresent); Sulawesi (Sulawesipresent) Australasia: Cape Province: present England: present Japan: present Madagascar: present NE. Madagascar: present New World: present New Zealand: present North Island of New Zealand: present Northern America:, Maryland (Marylandpresent) Old World: present Pacific: present Patagonia: present S. China: present South & Central Africa: present South Africa: present Southern America:, Peru (Perupresent) Tropical Africa: present U.S.A: present Ussuri: present W. Asia: absent W. Sumatra: present eastern U.S.S.R: present much of America: absent trans-Pacific: present
Four genera with about 80 species. Since VESTER'S (1940) small-scale map the family (Ascarina) has been found in Madagascar. It is mainly tropical but Ascarina extends south to North Island of New Zealand () and Chloranthus and Sarcandra extend north to Japan, China, Korea and the eastern U.S.S.R. (Ussuri).
Ascarina occurs in the Pacific and reaches New Guinea and the Philippines with a distinct section Madascarina in NE. Madagascar; Chloranthus and Sarcandra are widely distributed in Malesia, India, Indochina and China. Hedyosmum occurs in the New World from Mexico to Brazil and Peru and in the West Indies with one species occurring in the Old World in S. China, W. Sumatra, Borneo and Celebes ().
The family is now absent from Africa, W. Asia, Australia, and much of America.
HUMBERT & CAPURON (1955) when describing Ascarinopsis (= Ascarina sect. Madascarina) speak of it as part of the most ancient floristic element in Madagascar, a survival from the Cretaceous flora and AUBRÉVILLE (1976) considers it as an Australo-Papuan element similar to Hibbertia, Dillenia, Evodia, Protium (& America), Macadamia, Elaeocarpus, Weinmannia (& America), Bubbia, etc.
The complete absence of Chloranthaceae from tropical Africa at the present time is paralleled in many other groups with trans-Pacific distributions. The discovery of the fossil pollen type Clavatipollenites there could indicate that something very like Ascarina once occurred in Africa. The family might appear to owe its present distribution to a Gondwanaland origin in the Early Cretaceous or even earlier, the absence from Australia and Africa being attributed to climatic vicissitudes. Clavatipollenites (see later) is known from the Early Cretaceous of the U.S.A. (Maryland), England, Israel, Patagonia, South & Central Africa, Brazil, Australia, etc.; and probable Oligocene-Early Miocene deposits of South Africa (Cape Province) (COETZEE, 1981); and if all these refer to Ascarina or some closely allied genus then a different course of distribution is indicated. It seems the family may have been well distributed and common in the past but it is equally apparent that migrations involving any kind of stringent climatic deterioration are not feasible. Ascarina lucida for example was formerly (10,000-5,000 BP) abundant in New Zealand but is now much reduced due to increase of frost and drought (MCGLONE & MOAR, 1977).

Pollination

VAN DER HAMMEN & GONZÁLEZ (1960) have shown that Hedyosmum is wind-pollinated and has a high pollen production, but it has usually been assumed that the forest-dwelling Chloranthus and Sarcandra are insect-pollinated but I have traced no recorded observations. Some collectors mention scent.
Reference: .

Dispersal

The white-fruited Chloranthus erectus (= officinalis) is dispersed by birds according to RIDLEY () and the red-fruited Sarcandra glabra must also be.

Anatomy

This has been very thoroughly investigated by SWAMY (1953) and SWAMY & BAILEY (1950). An outstanding feature is the lack of vessels in the xylem in Sarcandra. The other genera have vessels but they are relatively unspecialised, Chloranthus being the least advanced. In Sarcandra tracheary elements in the secondary xylem are arranged in ± undisturbed radial sedations as seen in transverse section; tracheids in the region of the first year's growth measure nearly 1.9 mm and have very extensive overlapping ends similar to other vesselless dicotyledons indicating a cambium of very primitive type, and unusually long fusiform initials. The wood in Hedyosmum is of a very unspecialised type; parenchyma paratracheal as incomplete sheaths around the vessels; rays sometimes up to 1 mm wide, multiseriate rays composed of almost entirely upright cells; fibres with simple pits and occasionally septate wall rather thin; in Ascarina the parenchyma is apotracheal and the multiseriate rays of square to procumbent cells. Ascarina and Hedyosmum have nodes typically of the unilacunar type with 2 vascular strands in the leaf of Ascarina and 5 in Hedyosmum, the lateral pairs larger; Chloranthus and Sarcandra have modified unilacunar nodes with 5 vascular strands in the petiole, 2 much larger and extending most of the length of the midrib, the intermediate small trace disappearing about half-way but formed by the fusion of 2 minor branches of the larger traces at nodal level; the two small lateral strands come from a different gap. In Hedyosmum the stipular sheath formed from the connate petiole bases consists of collenchymatous tissue and supports the stem during intercalary growth. Lateral branches are initiated in the leaf-axils but are attached to the parent axis above the node at maturity; cork formed on the inner surface of the sheath brings it into intimate contact with the stem. There is a pulvinus on the stem at the upper margin of the sheath. In H. arborescens and related species the nodal sheath develops by pushing beyond the apical growing point and surrounding it, tightly closed above the bud and affording it protection. In Sarcandra and Chloranthus the stomata have 1 -2 subsidiary cells oriented parallel to the guard cells whilst in Ascarina and Hedyosmum there is a rosette of 4-6 ordinary epidermal cells.

BARANOVA (1983) reported that the laterocytic type of stomatal apparatus occurs in Chloranthus, Sarcandra and sometimes Hedyosmum along with other types. This type is known from a very heterogeneous mixture of families.

MELVILLE (1962) stated the 2 leaf traces in Ascarina can unite at various levels in the petiole or lamina and form a single vein, but in Chloranthus and Sarcandra each of the initial pair of traces forks, resulting in bundles, the middle pair reuniting to give a final triple trace. In Hedyosmum a trace of 5 bundles results from the bifurcation of the two outer bundles of such a triple trace. He points out these types are also to be found in both the Pteridosperms and Cordaitales.

Mucilage canals are present in the petiole, larger veins and also in the margins of the pith in Hedyosmum, in some species also containing sphaerocrystalline masses. Small clustered crystals are recorded in the inner part of the cortex. Stone cells are scattered in the cortex of younger stems of Sarcandra with larger groups in the pith; in older pith these cells form conspicuous transverse diaphragms alternating with plates of parenchymatic cells, but these diaphragms are absent from Chloranthus. Ethereal oil cells occur in the mesophyll of the leaf.

Taxonomy

Early placings of Chloranthus were various and mostly wide off the mark. LIND-LEY, who first recognised the family (1821) placed them in Piperales and his clear view has been supported by many authors (BENTHAM& HOOKER, 1880, 1883; ENGLER, 1894; ARMOUR, 1906; SWAMY, 1953; HOWARD, 1970; BEHNKE, 1975; BURGER, 1977) with arguments from various disciplines (morphology, anatomy, phytochemistry, pollen). After digesting a wholesale literature on the subject (1985), I believe this comes at least close to the correct affinity. The most probable conclusion from the floral structure appears to be that Chloranthaceae is a group of Piperales presenting in some points, especially in the structure of the ovary, primitive characters in common with the majority of the Archichlamydeae, while in other aspects modifications of the flowers are shown. A.C. SMITH (1972) found them more primitive than other elements of the Laurales and repeated (1981) that Chloranthaceae is best assigned to its own order, Chloranthales, validated by LEROY (1983). HUTCHINSON (1973) and CRONQUIST (1981) retain the family in Piperales, but this then remains a matter of choice.

It is certainly clear that the small, insignificant family Chloranthaceae is one of great importance in the study of primitive flowering plants whose ancestry points to high age, probably even to the Early Cretaceous, judging from the affinity of the pollen to Clavatipollenites. Unusual generic ranges also point towards early time, viz. one species of otherwise neotropical Hedyosmum in Indo-Malesia and one species of Ascarina in Madagascar even representing a separate section.

Cytology

EHRENDORFER et al. (1968) compiled lists of numbers for primitive families arid gave Hedyosmum arborescens n = 8; Sarcandra glabra 2n = 30; Ascarina rubricaulis 2n = 28; Chloranthus serratus n = 14&2n = 30; C. spicatus 2n = 30; C. japonicus 2n = 30; C. fortunei 2n = 60. He stated that from these data it can be seen that there are still true diploids in the family: ×= 8, further secondary polyploid base numbers ×2 = 14 and 15 and continuing infrageneric polyploidy (4×2). EHRENDORFER stated (in Beck, 1976) that "Chloranthaceae demonstrate progressive elimination of diploids (n = 8), major representation on the 4 × level (n = 14 and 15, the latter from 7 + 8?) and occasional origin of 8 ×(n = 30)", corresponding in part with the Piper ales.

Uses

The Chinese grow Chloranthus spicatus in pots for its fragrant aromatic leaves and it was formerly grown on a large scale in Java in one montane locality and used for imparting a scent to native tea and is apparently still used on a small scale for this purpose. BURKILL (1935) reported that leaves and flowers are left in contact with the tea whilst it is drying; they are later discarded or left to add bulk to the tea. Its use has been claimed to be deleterious. Among medicinal uses are as a poultice on carbuncles and boils, as a diaphoretic and excitant, as a cure for malaria but said to be poisonous in overdose, for back-ache and as a tea-like drink for treating coughs. Chloranthus erectus (= officinalis) is also used medicinally and as a tea, both the leaves and the roots being used. HEYNE (1927) gives extensive medicinal uses for the latter. It has a sudorific effect and is used in cases of fever and as a restorative during some phases of venereal disease; also as a stimulant and mixed with Cinnamomum bark as an antispasmodic during parturition.

DOSEDLA reported (GILLL. 1979) that formerly the leaves of Chloranthus erectus (= officinalis) were eaten together with pork as an offering in House Tambaran in the Hagen area of New Guinea. Areas where the plant grew were avoided; today the whole plant which stains the hands green on touching is taboo. VELDKAMP (in sched.) reported that in S. Borneo the branches are boiled and the slightly peppery tasting concoction drunk by women to prevent conception.

Sarcandra glabra has also been used to scent tea or add bulk and used in Indonesia for the same purposes as Chloranthus erectus. In the Philippines an infusion is said to be useful in treating headaches, asthma and 'internal pains'. It is also used in Chinese medicine for bone fractures, contusions and in the form of a leaf-decoction as an astringent treatment for vomiting. FRAKE reported (PNH 37989 & 38204) that on Mindanao the leaves are pounded and applied to ulcers and CHAI (S 35411) that the warm leaves are used for sprains. HEYNE also mentions that when tea-planting started in Java the cultivation of Chloranthus was forbidden by the Dutch.

Notes

The drawings on the four plates were made by Mrs. MAUREEN CHURCH, at Kew.

Phytochemo

Even to the collector the aromatic smell suggests relationship with the Piperaceae. In 1964 HEGNAUER complained that the chemistry of this family had scarcely been touched. BATE-SMITH (1962) found in the hydrolysed leaf-extract of Chloranthus erectus (= officinalis) none of the otherwise very widespread phenols save for β-coumaric acid and in this respect it resembles many Piperaceae. The ethereal oils which undoubtedly exist have so far not been chemically investigated.

KUBITZKI & REZNIK (1966) in their investigation of flavonoids as systematic characters investigated 4 species of Chloranthus, 1 of Hedyosmum and 5 of Ascarina and found them to contain traces to massive amounts of the two derivatives of flavanole, quercetin and kämpferol, the former dominating. SHIO& HIGUCHI (1981) have demonstrated the presence of a gymnosperm type of lignin, guaiacyl lignin in Sarcandra glabra and Chloranthus spicatus.