Illustrations of Indian Botany, Vol. 1/Anonaceae
This large, and truly tropical, order, is composed entirely of trees or shrubs, (no herbaceous member has as yet been found appertaining to it) with alternate, simple, entire, ex-stipulate, often fragrant, leaves ; and hermaphrodite, except in Hyalostemma, (Wall.) regular, axillary flowers. These are usually pale, or dull yellowish green, or brown, but sometimes yellow, and in Uvaria grandijlora are rich crimson, for the most part axillary, solitary, or a few together; but in Guatteria longifolia they form large clusters; in Artabotris, they spring from curiously hooked grapples, or tendrils, apparently abortive branches.
The calyx is generally deeply three parted; or rather perhaps, consists of three distinct sepals, partly united at the base; where they are very broad. The corolla consists of six petals, in a double series, three and three, often unequal; sometimes the exterior, sometimes the interior series, being much larger than the others; all caducous. The stamens are usually very numerous, sessile, and closely cover the whole of the enlarged, somewhat globose torus, rarely definite as in Bocagea : the anthers are two-celled, lateral, opening outwardly, and surmounted by an elongation of the connective, sometimes pointed, but oftener flattened, and truncated, more rarely they are adnate, as in Milium, a peculiarity which, when it occurs, promises to afford a useful generic distinction. The ovaries are generally numerous, one-celled, congested on the apex of the prominent torus, either free or united : ovules few or numerous, variously attached; being either numerous and transverse, springing in a double row from the inner angle of the cell in Uvaria; or solitary, and erect, and from the bottom of the cell in Guatteria; or pendulous from the top of the cell in Orophea. Style usually short or wanting, stigma capitate. The fruit is apparently more variable in its character than the other parts of the fructification, and has hitherto been almost entirely looked to for generic distinctions. In some, it consists of numerous, united, one-seeded, carpels, enveloped in soft pulp, and forming together a pulpy fruit, as in Anona; (the custard-apple, sour sop, and bullock's heart) in others, the cai-pels have one or several seeds, and are borne on along peduncle, as in Uvaria, Guatteria, Sec. and sometimes these carpels though otherwise distinct, remain sessile, or with the peduncles so short that the carpels form together a globose head : (this variation occurs in some species of Uvaria, and Miliusa.) In others, as Unona, the carpels are elongated, containing several seeds, and contracted between them like a necklace of beads. The seeds universally possess the remarkable character of having ruminated albumen, like those of the nutmeg. A few have them arilled as in that genus, which (aril) when it exists, is supposed to secrete at the base of the seeds, a resinous highly aromatic matter. This is the case in what is called the Ethiopian pepper, ( Habzelia Ethiopica J and some others, which, partly on that account, the younger DeCandolle has united to form the genus Habzelia, a genus common to both Africa, and America, but not yet found in Asia.
Affinities. The affinities of Anonacece are so various, as not to be easily indicated in a few words, but their closest alliance is certainly with Magnoliacece, from which however, they are readily distinguished, by their ex-stipulate leaves, their more distinctly formed, and sub-persistent calyx, by the form of their anthers, the arrangement of their ovaries, but above all by their ruminated albumen, This last character however, combined with the ternary arrangement of their flowers, the occasional presence of an aril, and their aromatic properties, so closely associates them with Myrslicacece, (the nutmeg tribe) that Professor Lindley seems to consider the latter, as little ehe than an apetalous form of Anonacece. Thus constituted, it is difficult to give an abridged character of the order, but as, generally speaking, the Indian species present the normal forms, their character may be summed up in the following terms.
Essential Character. Polypetalous, polyandrous, ovaries wholly superior : carpels more or less distinct : very rarely solitary : seeds, usually without an aril, albumen aromatic, ruminated. Leaves alternate, ex-stipulate. In one Indian genus ( Hyalostemma J the flowers are diaecious.
Geographical Distribution. This, as already remarked, is strictly speaking, a tropical order, confined to Asia, Africa, and America, none having as yet been found in Europe, or Australia, and very few beyond the 30th degree of latitude. In 1832. when Alphonse DeCandolle published his memoir on this family, he reckoned 204 as the total number of species, of these 87 are Asiatic, 95 American, and 22 or 23 African. He thinks, however, that the relative proportion of the Anonacece, to the total vegetation, is nearly the same in all equatorial regions. In 1818, the number of known species amounted to 105, hence in the short interval of 14 years, the number of species was doubled, and several have since been added. Dr. Wallich enumerates about 80 Indian species, to these one or two were added by Mr. Arnott and myself, and several other peninsular species, have been recently discovered in the course of my excursions : doubtless, many yet remain in our jungles, especially in those tracts of hilly country, enjoying a moist climate, with a moderate range of temperature. From this I infer, that when such localities have been more carefully examined, the Indian list will be found to exceed 100 species; which I consider the more probable, as I observed at Courtallum, where I found the greatest number, that they always occupied the thickest woods, and being themselves, generally of a very plain, and unostentatious aspect, with small flowers, are apt to be passed unnoticed. In proof of the correctness of this opinion, I believe it is only necessary to add, that when DeCandolle and Dunal wrote, the number of known, Indian species, amounted to only thirteen.
Properties and Uses. Like most other tropical orders, the properties and uses of Anonacece, are imperfectly known to Europeans; and in this country at least, a very few have as yet been admitted into cultivation, though some of them are sufficiently ornamental. For their fruit, three species of Anona, are cultivated in India, A. squamosa, (custard-apple) A. reticulata, (bullock's heart) and A. muricata, (sour sop). This last I have not myself seen in India, but being the A. asiatica of Linnaeus, must have been one of the first introduced.
The Guatteria longifolia (or Asochum) is commonly cultivated about Madras, and some other places, as an ornamental tree; a distinction to which its elegant form justly entitles it, but its wood is said to be soft, and of little use. Artabotris ordoratissima, a diffuse shrub, with exceedingly fragrant flowers, springing from a curious hook, or grapple, is occasionally, though too rarely, cultivated as an ornamental shrub, which it well merits. These, to the best of my recollection, are the only members of this order, I have met with in cultivation. Many others both of Indian, and Exotic origin, "might however be brought, with advantage, into our gardens. The Uvaria grand/flora, a superb plant, a native both of Sumatra, and Ceylon, which has succeeded well in the Calcutta botanic garden, has large showy bright crimson flowers. Uvaria narum the one represented here, might also form a desirable addition to the few ornamental shrubs, we possess. Guatteria virgata, (the lancewood of Jamaica) the wood of which is highly esteemed in England, on account of its strength, and elasticity, might form a very useful addition to our stock of woods, suited for the operations of the coachmaker.
Most species of Anonacece are remarkable for their fragrance; and for their aromatic properties, diffused through every part of the plant. The roots of the Uvaria narum enjoy these properties in a high degree; and are, we are informed by Rheede, much employed in Malabar, in the cure of a variety of diseases. He states that the infusion of them, is successfully administered in some forms of fever, and hepatic disease; that bruised in saltwater and rubbed on the skin, they cure the morbus pedicularis, originating in general weakness. This property of destroying vermin, is equally enjoyed by the seeds of the custard- apple, which, we are informed, by Mr. Royle, are, in Bengal, powdered and mixed with flour of Bengal gram, ( Cicer Arietinum J and used to wash the hair for the purpose of destroying these unwelcome intruders. Contrary to the usual character of the tribe, the leaves of the custard-apple have a heavy disagreeable smell. Upon the whole I think it may be safely asserted, that rich as India is, in species of this order, we as yet know very little of their useful properties, and judging from analogy, it may with equal safety be asserted, that they should be carefully inquired into. For example, we already know that most of the species, of Habzelia, natives of both Africa, and America, are highly aromatic, and that the seed of one, or more of them, forms a considerable article of commerce, under the name of Ethiopian pepper. In Xylopia, (Bitter wood, so called from the taste of the wood) a West Indian genus, the bitter principle so universally prevails, that every part of the plants possess it. In X. glabra " the wood, bark, and berries, have an agreeable bitter taste, not unlike that of the orange seed. The wild pigeons feed much on the latter, and owe that delicate bitterish flavour, so peculiar to them in the season, wholly to this part of their food. Fresh gathered from the tree, they are agreeable to the palate, and grateful to the stomach.
The bark is also richly impregnated with this juice, as well as the wood, and both yield a very agreeable bitter in the mouth while fresh ; but. that diminishes greatly after they are dried. The bitter quality of this tree is communicated with great facility. A handful of the shavings immersed in water, and instantly taken out again, will render it of a very bitter taste. Sugar sent over in hogsheads made of this wood, was so bitter that no person would purchase it. Bedsteads, and presses made of it are proof against cockroaches and other insects."* A decoction of it is said to create appetite, and possess the usual tonic properties of bitters. These examples are I presume, sufficient to establish the value of the properties inherent in members of this family, and go far to prove, that among the numerous Indian species belonging to it, we have good reason to expect, some, when properly investigated, will be found not less valuable,, than the better known American and African ones.
Remarks on the Genera, &c. Of this order Dr. Roxburgh, in his Flora Indica, gives characters of 27 species, classed under three genera, namely, Anona, Uvaria, and Unona ; Guatteria, was afterwards added to the catalogue, of Indian genera, by referring several of Roxburgh's Uvaria's to that genus. These genera were distinguished by characters almost entirely taken from the fruit, without reference to the ovary, or indeed to any other part : Anona, being distinguished by having a number of carpels, united into a single compound fruit: Uvaria, by having its carpels distinct, stipitate, fleshy, many-celled, and many seeded ; with the seeds ranged in two rows : Unona, by having distinct, but dry, many-seeded, carpels ; the seeds ranged in a single row, and often resembling, by the contraction of the carpel between them, a necklace : Guatteria, by having dry, globose, stipitate, one-seeded, carpels. These simple, and, at first sight, apparently all-sufficient characters, were found on more careful examination to be exceedingly incorrect. Uvaria, for example, to which was attributed a many-celled fruit, and two rows of seed, was found to have a one-celled ovary, and the rows of ovules, if two, so close as scarcely to be distinguishable, and in truth forming a single line attached to the inner angle ; hence the many cells, and two rows of seed observed in the mature carpel, must be produced by mere condensation of the surrounding pulp, and the divergence of the free extremities, of the seed, since they are all attached along the same angle of the seed vessel. In Unona, the structure of the ovary is the same, with probably fewer ovules, hence it follows, that the abortion of a few ovules (by leaving more room for the regular development of the remainder) may convert a Uvaria,'mto&Unona; and vice versa, an unusual number of ovules, or any hindrance to the usual course of development of the seed vessel, might equally change Unona, into Uvaria ; the differences between the two genera, thus rest on adventitious, not structural, differences. In support of the justice of this position, it is only necessary to state, that a large proportion of the species of the latter genus, have, since the publication of DeCandolle's sy sterna, been removed to the former. The propriety therefore, nay, the necessity, of uniting the species of both, and of two American genera, having similar ovaries and fruit, (Asimia and Porcilia) into one genus, as ably advocated by M. Richard, in some remarks on the subject in the Flore Senegambie, becomes evident. Blume, however, as appears from DeCandolle's memoir, has revised the character of Uvaria, and still keeps them distinct ; but as I have not his work to consult, I am unable to state with what propriety. Guatteria, which in like manner is characterized from the mature fruit, without reference to the ovary, may be simulated by species of Uvaria, or Unona, through the abortion of all the ovules but one, a modification of which my collection presents specimens.
Swayed by these facts, M. Richard proposes an amended character for Uvaria, in which the one- celled, many ovuled ovaries, with the ovules attached along the inner angle, forms the essential distinction; a modification which admits of the association of all the species now referred, to the four genera above named. The character of Guatteria, might be similarly modified with advantage, and would then, perhaps, be found to separate the American, from the Indian, division of the genus. All the Indian ones I have yet examined have a single, erect, ovule, attached to the bottom of the ovary. Whether or not the American species referred to this genus, possess this structure, I am unable to say, but in the following Indian ones, I have ascertained it to exist. Guatteria longifolia, Korinti, cerasoides, and suberosa. By this addition to the character, all Anonace(s.vth single seeded carpels, but having more than one ovule, or even with one ovule pendulous, or attached to the inner angle, and placed transversely across the cell of the ovary, will be excluded. In combination with this structure of the ovary, I have in all the above instances, found the petals nearly equal, and the connectivum of the stamens truncated, that is, prolonged beyond the anthers, and ending in a broad shield-like apex.
Four other genera have more recently been added, to the peninsular flora, jtrtobotris, Brown, Milium, Alph. DeCandolle, Lobocarpus, Wight and Arnott, and Orophea, Blume. The first of these is well known: the second is described, and figured, by the younger DeCandolle, in a memoir in the Geneva, Natural History Society's, Memoirs, but apparently from very imperfect specimens, since his character differs in some essential points, from numerous specimens, I have collected since my return to India, and most of them from the same station, from •which his was procured, namely, Courtallum. DeCandolle assigns to his, 3 sepals, and 3 petals, united half their length, and forming a bell-shaped corolla, at the base, that is two series of verticels. In mine, there are invariably three verticels, the two outer ones much smaller, both hairy, and sepaloid, in aspect, giving it the appearance of having 6, in place of 3 sepals, and a 3-petaled corolla: while the stamens are represented by DeCandolle as only 12 in number, and placed in a single series, round the elevated torus on which the ovaries are borne,while in mine, I have always found them, as in most of the other members of the order, numerous, closely covering the whole of the torus, and nearly sessile, in place of being furnished with a distinct longish filament. I think I have now two species of this genus, one of which, in spite of these discrepancies, I consider the same as DeCandolle's. Lobocarpus, I have not again met with, and have nothing to add, to what is contained in my flora. Orophea, which I lately found at Courtallum, in dense thickets, is a handsome shrub, but with small inconspicuous flowers. It is readily distinguished by having the inner, series of petals, much larger than the outer, and attenuated at the base, into a distinct claw or unquis. The stamens are in a double ? series about 15, and have the appearance of being 4-celled, from a depression along the line of dehiscence. Ovaries with 2 pendulous ovules, and furnished with a large style. Some other, perhaps new genera and species, of which I have specimens, remain to be added, but as I have not yet been able to ascertain, whether or not they are already named, I forbear for the present, characterizing them, as it is my intention, so soon as leisure permits, to re-examine the whole series, and figure, and describe all those that are new, or but imperfectly known. One however, of these genera, of which I possess two species, both from Ceylon, distinguished by some unusual peculiarities, is too interesting to be passed over in silence on the present occasion.
In the form of the corolla and torus, this genus associates with Ccelocline, but differs so widely in the character of its ovaries and fruit, as leaves me no room to doubt its being new and perfectly distinct. The most remarkable peculiarities observed in the course of my, perhaps too hurried, examination, were—1st. The calyx is persistent, the sepals united to near the apex, enlarging with, and enclosing, the fruit.—2d. The torus is concave, prolonged beyond the ovaries, bearing the stamens exteriorly, and the ovaries within.—3d. The ovaries are indefinite, few (one ?) ovuled,with the ovules attached near the apex, and pendulous.—4th. The styles are long, subulate, ending in a simple acute stigma.—5th. The carpels are one-seeded, united into a single fruit, enclosed within the enlarged calyx, and ranged in a single row, round the central axis, with the solitary compressed seeds, pendulous from the apex. The fruit, in my specimens, is not quite mature, but is sufficiently advanced to enable me to ascertain these points. This genus may therefore be thus characterized.
Patonia, R. W.
Sepals 3, united at the base, persistent, enlarging with the fruit. Petals, in a double series, distinct. Stamens numerous, distinct, truncated at the apex. Torus concave, bearing stamens externally, and ovaries within. Ovaries free: ovules solitary, (?) pendulous from the apex. Style longish, subulate. Stigma acute. Carpels numerous, ( 10-15) all united, enclosed within the enlarged tubular calyx. Seeds solitary, compressed, pendulous from the apex of the carpel. Trees or shrubs : leaves oblong, lanciolate, acuminated, glabrous : flowers axillary, solitary, or several together, from a short peduncle, or abortive branch: corolla slender ; petals tapering towards the point, pubescent : fruit, concealed within the tube of the greatly enlarged, 3-lobed calyx.This genus is dedicated to Miss Paton, now Mrs. Colonel Walker. The name Walkeria being pre-occupied, prevents me dedicating it to my highly esteemed and amiable friend, under the name by which only she is known to science, as the diligent and acute investiga tor, and, not less graphic than persevering, delineator, of the plants of Ceylon.
1. P. Walkerii (R. V.) Leaves glabrous, oblong-lanciolate, acuminated : lobes of the calyx, acute in flower, obtuse in fruit : corolla somewhat ventricose. — Hab. Ceylon, in woods: communicated by Colonel Walker.
The leaves are about 6 inches long, by 2 broad, with a long acumen, shortly attenuated at the base, glabrous on both sides. The corolla is somewhat ventricose in the middle, contracted within the calyx, and towards the point; lobes of the calyx very acute, and reach- ing to the dilated portion of the corolla, as the fruit advances, becoming broad, obtuse, and reflexed at the point.
2. P. parvifolia (R. W.) Leaves lanciolate, acuminated, glabrous : lobes of the calyx broad, very obtuse, hairy : corolla gibbous at the base, attenuated upwards, tomentose : fruit —
Hab. Ceylon, communicated by Colonel Walker.
A shrub with glabrous, lanciolate, acuminated, leaves ; from 1 1 to 2 inches long, and about I broad. Corolla very gibbous, or sub-globose, at the base, and thence attenuated to a point. Calyx lobes, broad, obtuse, shorter than the dilated portion of the corolla. The fruit of this species is still unknown, hence it may possibly prove a species of Caelocline.
EXPLANATION OF PLATE 6.
1. Flowering branch of Uvaria Narum.— 4. Peduncle, and cluster of full-grown, stalked, carpels. — Natural size.
2. Sepals, stamens, and ovaries.
3. Flower, cut vertically, showing the ovaries and stamens on the prominent torus.
5, 6. Back and front view of the stamens. — 7. Ovary detached. — 8, 9. The same cut transversely, and longitudinally, showing the position of the ovaries. — 10, 11. A carpel cut transversely, and longitudinally, showing the position of the seeds, and the spurious cells of the fruit. — 12. A seed. — 13. The same cut vertically, to show the ruminated albumen— all more cr less magnified.
- * Loudon's Encyclopaedia of Plants. Art. Xylopia.