Illustrations of Indian Botany, Vol. 1/Xanthoxylaceae

by Robert Wight

XLIX.—XANTHOXYLACEAE.

As stated above, this is viewed by most Botanists as merely a section or suborder of Rulaceae, but is certainly an interesting one, including many genera, yet, all so intimately united that it becomes exceedingly difficult to determine their limits. Though in a great measure of tropical origin the Peninsular flora embraces but few representatives, our list in the Prodromus only extending^, including Ailanthus, to seven species ranged under three genera, namely, Xan- ihoxylum, Toddalia and Ai.lant.hus, Mergui and Malacca have Brucea in addition. One addi- tional genus may however be required for the Peninsular species. Generally the species are either trees or shrubs, sometimes scandent, frequently armed with strong prickles, having, usually, compound leaves pierced with numerous transparent oily glands, like thos3 of Aurantiaceae, and numerous aggregated small, generally, unisexual flowers : that is, male flowers furnished

with perfect stamens and the rudiments only of an ovary, or with a perfect ovary and imperfect stamens. Ailantheae are usually united with these as a mere tribe, Dr. Arnott in my opinion more judiciously forms of them a suhorder, an arrangement, which I adopt here, hut for the purpose of exhibiting both views I subjoin in his own words Meisner's characters of these tribes* as given " in aureo CI. Meisneri libro cui titulus: Plantarum Vascular inm Genera tabulis diagnosticis exposita" a work well meriting the brief but forcible encomium thus bestowed by the celebrated DeCandolle, himself, the author of the noblest Botanical work that has yet issued from the press, a monument of the most untiring industry, and most profound research.

"Flowers by abortion unisexual, regular. Calyx 3-4-5 divided. Petals equal in number (rarely more) to the sepals : aestivation usually twisted, convolute. Stamens as many, or twice as many, as the petals, inserted round the base of the torus. Torus elevated and forming a gynophore or short thick stalk to the pistillum, which, in the male flowers, is rudimentary or rarely entirely absent. Carpels usually as many as the petals, sometimps fewer, seated on the gynophore, sometimes combined into one ovary, sometimes entirely or partially distinct* ovules 2, or rarely 4, in each carpel: styles in the single ovaries combined, in the distinct ovaries either distinct or combined upwards, sometimes none : stigma 2-5 lobed in the united styles, simple in the distinct styles. Fruit sometimes single, baccate or membranaceous. 2 5 celled : sometimes of 1-5 distinct drupes or 2 valved capsules, of which the sarcocarp is either entirely combined with, or only partially separable from, the endocarp. Seeds solitary or in pairs, pendulous. Embryo lying within a fleshy albumen: radicle superior: cotyledons oval, flat. — • Leaves exslipulate, alternate or opposite, with pellucid dots or rarely without them."

Affinities. These are so complex that I abstain from attempting to elucidate them myself, and therefore have course to the far abler exposition given by the younger Jussieu, and reprinted in Lindley's Natural System of Botany, merely observing, that I am not disposed to lay so much stress on characters taken from the absence or presence of albumen in the seed, since every day's observation tends to convince me that characters taken from it require to be used with caution, as I think our knowledge of vegetable structure, in connection with the absence or presence of this substance in the seed is not sufficiently advanced to enable us to

Tribus I. XANTHoxLEa; — Arn.

• Genuinse : florib-diclinib. (rarissime liermapbrod. cfr Picrasma).

A. Discus elevatus, gynophorum breve crassum forrnans. Ovaiia in fi. £ rudimentaria, rarissime Q, in fl. $ plus minus inter se coalita, rarius distincta rarissime (v. Blackburnia) unicum, singula 2-rarius 4-ovulata. Stig- mata distincta simplicia, aut in unicum 2-5-lobum connata. Carpella 1-5 disiincia aut coalita, nunc indebisceniia drupacea, nunc capsularia 2-valvia. Albumen carnosum. Embryo saepius curvatus. (Folia nonnunquam oppo- sita, plerumque pellucido-punctata.)

Tribus II. AilanthejE— Arn. Discus depressus aut cupulwformis, 4 lobus. Ovaiia plura, distincta, 1-ovulata. Stigm. distincta. Carpella 3-5, distincta, indehisc, drupacea aut samaroidea. Albumen ? tenue, carnosum, seminis integumento adheerens. Embryo rectus. (Folia altema, impunctata. Flor. ssepe polygami.)

Xanthoxylum. (L.) Kth.

1 Ovula juxtaposita. Cal. 4-5-part. Pet. 4-5, rarissime O. 9 : Stam. sterilia squamiformia, ananthera aut anth. effoetas gerentia, aut O- Ovaria 5-1. Styli liberi aut apice connati, interdum subnulli. Stigmata libera capitata, aut inter se cobaerentia demum solubilia, aut (sicut styli) in unicum 4-5 lobum connata. Caps. 5-1, sessil. "V. stipitatse, distinctae, rarius inius connaiee, 2-1-spermse. (Arb. V. i'rut., segpe aculeat. Fol, simpl. V. compos, pell.-punct. Inflor. varia. — Patria extra Euroji. diffusa, pra?cipue intra trop.

• Flores apetali, partium numero quinario. Sepala petaloidea (ex K until interdum) Xanthoxidon Colden

9). Styli stigmate clavato conjuiicti, basi distincti $ - J

nt. Pet. 3. Germ. 3-part. Styli 3, fili- )„,,.. n , n n ■> n f f a A t J ovinia Desv.

. Carpella 6, 1-spenna.— (Ins. Antiu .).. )

6

• • — —Completi.

A. — Partium numerus ternarius. Cal. 3-dent

form

B. qualernarius.

1) Ovaria 2 Pterota. P. Brown

2)

g T—. 4 -. %li4,stigmatib.. I Auhertia. Bory. S. Fine

Capilellatis inter se colieerentes *

quinarius. Cal. 5^ part. Pet. 5. Stam. 5

1) Ovaiia 3-5 Ochroxylum. Schreb.

2) 1 , Lanysdorfia. Leand. draw useful characters from it. Of the genus Zanthoxylon for example, in all the species I have examined, 4 in number, the cotyledons are large with the albumen, if indeed such it be, reduced to a mere membranous covering. In Tuddalia bilocularis it is altogether wanting. True these may, and perhaps ought to be removed from the genera to which they are referred on that very account, but the fact of its absence in some and presence in others, shows how value- less it is an ordinal character in this tribe, and is still further shown by what is observed in Rutece and Diosmece, which, though so nearly related, are yet distinguished by the one tribe having albumen, the other being exalbuminous. Having premised these few remarks on this point of structure, I shall proceed with the extract.

"This is one of the families which comprehend genera with both distinct and concrete carpella; the latter are often entirely distinct, even in the ovarium, but more frequently there is a union, or at least a cohesion, of the styles, by which their tendency to concretion may be recognized. In a few instances the carpella are absolutely solitary.

"The place originally assigned, and for a long time preserved, for most of the genera of Zanthoxyleae, proves sufficiently how near the affinity is between them and Terebinthaceae. If, with Messrs. Brown and Kunth, the latter are divided into several orders, Zanthoxyleae will be most immediately allied to Burssraceae and Connaraceae, agreeing with the former in the genera with a simple fruit, and with the latter in those with a compound one. Notwithstanding the distance which usually intervenes in classifications between Aurantiace.ae and Terebinthaceae, there are nevertheless many points of resemblance between them ; Correa has pointed out a passage from one to the other through Cookia, Kunth, in new-modelling the genus Amyris, and in considering it the type of a distinct order, suspects its near affinity with Aurantiaceae ; we cannot therefore, be surprised at the existence also of relations between the latter and Zanthoxyleae. A mixture of bitter and aromitic principles, the presence of receptacles of oil that are scattered over every part, which give a pellucid dotted appearance to the leaves, and which cover the rind of the fruit with opaque spaces, — all these characters give the two families a considerable degree of analogy. This has already been indicated by M. de Jussieu in speaking of Toddalia, and in his remarks upon the families of Aurantiaceae and Terebinthaceae; and it is confirmed by the continual mixture, in all large herbaria, of unexamined plants of Terebinthaceae, Zanthoxyleae, and Aurantiaceae. The fruit of the latter is, however, extremely different; their seeds resembling, as they do, Terebinthaceae, are on that very account at variance with Zanthoxyleae, but at. the same time establish a further point of affinity between them and some Rutaceous plants which are destitute of albumen. Unisexual flowers, fruit separating into distinct cocci, seeds solitary or twin in these cocci, enclosing a usually smooth and blackish integument, which is even sometimes hollowed out on its inner edge, a fleshy albumen surrounding an embryo the radicle of which is superior, are all points of analogy between Zanthoxyleae and Euphorbiaceae, particularly between those which have in their male flowers from 4 to 8 stamens inserted round the rudiment of a pistil, and in the female flowers cells with 2 suspended, usually collateral, ovules. Finally, several Zanthox yleous plants have in their habit, and especially in their foliage, a marked resemblance to the ash. The dioecious flowers of Fraximm, its ovarium, the two cells of which are compressed, having a single style, 2 ovules in the inside, and scales on the outside, and which finally changes into a samara which is 1 -celled and I -seeded by abortion, all establish certain points of contact between Ptelea and Fraximus." Adde Juss.

Geographical Distribution. The greatest number of the plants of this order are found in tropical America, a few are natives of Africa: two of which, Zanthoxylon Senegalense and Z. Leprieurii resembling our Z. Rhetsa, are from Senegambia. On continental India the number hitherto discovered is not great : they appear more numerous in the Islands to the eastward. Blume, including Rutea, has from Java 14 species, while Roxburgh has for India only seven, Wallich's list has about 28 for all India. The Peninsular flora at the time of our publication only presented a catalogue of 10 species for the whole order Rutacece, but for Zanthoxyhce excluding Rutece and AHanthece only 5, one of which is since excluded, and the genus of another still doubtful. In addition to the species described in the Prodromus, I have since found Zanthoxylon (Fagara, Roxb.) triphyllum and two new species referable to the subgenus Langadorfia, also Toddalia Jloribunda, Wallich, thus adding four species to our list which now exceeds Roxburgh's. Properties and Uses. Bitter stimulating and aromatic properties pervade in greater or less intensity, almost every species of the order, at least so far as our acquaintance with them yet extends. Some species are remarkable for their tonic properties, others are powerful sudorifics, and when applied to the gums or even taken internally act as nowerful sialogogues.

These properties point them out as suitable remedies for Rheumatism, in which complaint they have been found very useful. Z. alatum of Roxburgh, a Nepwul plant, is aromatic and pungent, and the seeds are used medicinally by the natives. Another species, Z. piperitum, a native of Japan, but now cultivated in India, is a powerful aromatic and used in its native country in place of pepper. It is also used as rubifacient and discutient by the natives, being applied in form of a poultice to the neighbourhood of inflamed parts. The seeds of Z. Budrunga, (Roxb.) have the fragrance of Lemon peel, and being of a warm spicy nature are used medicinally by the natives of Silhet, where it is indigenous. The unripe capsules Z. Rhetsa are like small berries and are gratefully aromatic, tasting like the peel of a fresh orange. " The ripe seed taste exactly like black pepper but weaker, from this circumstance I conceive this may be F. perperita, yet I have always found the leaflets entire" (Roxb.) The inner lamina of the bark is also acrid and bitter. Of Toddalia aculeata, a very common plant in this country, Roxburgh remarks Fl. Ind, 1 page 617 — " Every part of this shrub has a strong pungent taste. The roots when fresh cut smell particularly so. The fresh leaves are eaten raw for pains in the bowels; the ripe berries are fully as pungent as black pepper, and with nearly the same kind of pungency ; they are pickled by the natives, and a most excellent one they make.

"The fresh bark of the root is administered by the Telinga physicians for the cure of that sort of remittent, commonly called the hill fever. I conceive every part of this plant to be possessed of strong, stimulating powers, and have no doubt but under proper management it might prove a valuable medicine where stimulants are required."

Remarks on Genera and Species. The number of genera of this suborder is not considerable, amounting in all, according to Lindley, to 15, including Ailantherz. Of these Zanthoxylon seems to have been the most troublesome to Botanists, there being no fewer than seven generic names ranged under it as synonyms — whether all these ought to be reduced I am not prepared to say, but, judging from what I have seen in examining the few species in my collection, I suspect some at least of these will be restored, or if they are not, then, on the same principle, 1 think many genera which now encumber the Botanical system might be similarly reduced.

For the purpose of facilitating the investigation of this polymorphous genus I have added to the proceeding extract from Meissner's work the characters of the subgenera of Zanthoxylon. Comprehensive however as the character of this genus is our Z. cannarioides must be removed. In it there are 5 sepals, 5 petals, 10 stamens, alternately longer and shorter, the filaments united, forming a tube, hairy within, enclosing the style, and large stigma, inserted, with the petals, outside of a large cup-shaped disk in which the 2- celled ovary is immersed, seed exalbuminous, with the radicle next the hilum. These peculiarities of the flower com- bined with the exalbuminous seed afford ample reasons for its separation, not only from the genus but from the order : in a word, it is a species of Heynea. Our Toddalia bilocularis must also, I fear, be equally removed from that genus, at least, if the absence of albumen, or its presence in a very unusual form, can be received as a character of sufficient weight, when aided by the difference of structure of the seed itself. The flowers however are still unknown, and unless they assisted in distinguishing this as a distinct genus, I should hesitate in removing it from the place it now occupies on account the peculiarities of the fruit only. Besides wanting albumen, the seed are remarkable for the extreme inequality of their cotyledons, the one, being three or four times larger than the other, 1 have not yet found flowers but judg- ing from the seed, I should suppose it more nearly allied to Aurantiacece than Zanthoxylacece, and certainly forms an additional link between these in many other respects allied orders. Between Evodia, Blume, and Fagara, Roxburgh, there appears to be no difference, and indeed between Evodia, Juss. and Zanihoxylon, (subgenus Aubertia) the only difference seems to be in the declinous flowers of the latter. This distinction can scarcely be considered a good one, though on such only does the essential distinction between Rutece and Zanthoxy/ecs rest, because, the rudiments of all the organs of perfect flowers constantly exist in both sexes, and only require a slight change of circumstances to render them perfect : on this account, I sus

pect both Rlume and Roxburgh bave described the flowers of their respective genera, Evodia and Fagara, erroneously in making them perfect and bisexual. Of the former I cannot be certain, but can have no hesitation with reference to the latter Botanisl having now before me male flowers of his F. Rhetsa and F. triphylla, the ovary of which, though apparently perfect, yet wants ovules and is sterile. Such is probably also the case with all Blume's species of Evodia, which must in that case, be brought here. This is the more likely to be the case, as all the other species of Evodia are from New Holland.

ZANTHOXYLON.

Z . triphyllum : (Juss.) Unarmed, leaves opposite, palmately trifoliolate, leaflets oval, oblong, acuminated, somewhat unequal sided at the base, glabrous: peduncles axillary, longer than ti e petiols, corymbs large, spreading; flowers numerous, minute: carpels 1-4, spreading, one seeded. Seed globose, glossy black.

Fagar triphylla, Roxb. Evodia triphylla, D'C.

Z. (Langsdorfia) ovalifolium , R. W. Unarmed, shrub- by, leaves alternate, palmately 3 foliolate, leaflets oval, shortly acuminated, somewhat cuniate at the base, entire, glabrous on both sides: panicles axillary, longer than the leaves, contacted, the short side branches ending in small capitate clusters of flowers. Fruit about the size of a pea, 1-seeded. Seed globose, cotyledons thick and fleshv, .dbuinen thin, membranous.

Sh^a'jerry kills in flower, and fruit in August and September.

Z. (Langsdoifia) sepearinm — Rigid, ramous, armed with numerous straight prickles: leaves palmately 3

foliolate, short petioled, leaflets oval, oblong, obtusely acuminated, crenulate on the margin, coriaceous, gla- brous : panicles shorter than the leaves, contracted, speciibrm, few flowered. Fruit about the size of a pea, 1-seeded. Pulicat hills near Madras.

This species is evidently very closely allied to the preceding, but is yet so different in every part of its habit that I cannot think of uniting them.

Toddalea aculeata and T. floribunda. In our Prodromus we remark that between the speci- mens of my catalogue No. 532, and Wallich's T. flori- bunila,v/e could point out no character to separate them, unless the tubercles on the stem. The plant here figured is identical with the specimen alluded to, and has, as I have since found, the tubercled stem represented in Wallich's plate ; hence is identical with it also. I therefore think (hat this may be received as a good species, and beg that the name'on the plate may be changed from aculeata to floribunda.

EXPLANATION OF PLATE 66.

1. Toddalia floribunda, Wallich — natural size.

2. An expanded flower.

3. Stamens.

4-5. Sections of ovaries, showing them both sterile, and that this is actually a unisexual genus.

6. A female flower.

7. The ovary, the petals removed.

8. cut transversely, 5-celled.

9. cut vertically, ovules 2 in each cell,

superposed.

10. A young fruit.

11. The same cut transversely.

1 2. A seed — natural size.

13. magnified.

14. cut transversely, showing the cotyledons

enclosed in a thin albumen.

The figures representing the female flower were taken from specimens of T. aculeata, those of the other being all male plants.

SUB-ORDER—AILANTHEAE.

This sub-order has been in a great measure treated of under the preceding, little therefore remains for this place. The genera referred to it were formerly placed by DeCandolle in his tribe Connaracece of the order Terebinthacece. Farther examination has led to that, at the time he wrote, ill understood order being greatly modified and to the removal of several of his tribes and genera to form new and distinct families. Those referred here were then transferred to Rutaceae, and more recently were by Dr, Arnott established as a 2d sub-order of that extensive order, an arrangement, partially adopted by Meissner who considers them a distinct tribe of Zanthoxylaceae. They principally differ from their associates in having solitary ovules and drupacious or samaroid (winged) fruit. Dr. Arnott gives the following character.

Sub-order 2.— Jlilantheae (Am.) Flowers unisexual, regular. Calyx 4-5, divided. Petals 4-5, alternate with the sepals : aestivation between valvular and twisted. Stamens as many as the petals and alternating, or twice as many, about the same length: filaments inserted round the base of the torus, not arising from scales : anthers 2-celled, bursting longitudinally. Torus discoid, or 4-lobed and cup-shaped. Ovaries several, distinct : ovules solitary in each cell, pendulous, styles very short, distinct, or slightly cohering at the origin of the stigmas : stigmas (distinct, filiform, recurved, papillose. Carpels 3-5, distinct, indehiscent, drupacious or samaroid, 1-celled, 1 -seeded. Albumen ? a thin fleshy plate, closely adhering to the integument of the seed, and resembling an inner coat. Embryo straight : radicle superior, short : cotyledons fleshy. — Trees or shrubs. Leaves alternate, exstipulate, equally or unequally pinnated, without pellucid dots.

Affinities. The affinities of this tribe have been already sufficiently explained.

Geographtgal Distribution. All the species of Ailanthus are of Asiatic origin, two are natives of India, one of the Moluccas and one of China. Of Brucea the original species was found by Bruce in Africa, to whom the genus is dedicated, one is a native of India, and one of Sumatra and the Malayan peninsula, of which I have also specimens from Ceylon, but according to Moon introduced. Eurycoma, a genus doubtfully referred here, is a Malayan plant.

Properties and Uses. The bark of Ailanthus excelsa £f has a pleasant and slightly

bitter taste, and is prescribed by the native practitioners in decoction in cases of Dyspepsia"

Ainslie. In confirmation of that statement I may add that some time ago specimens of this plant were sent me, as those of a tree, the bark of which is prescribed in the Circars as a pow- erful febrifuge, and tonic in diseases of debility. The wood of this species is white and soft, that of A. glandulosa, the Chinese species, is hard, heavy, glossy, like Satin, and is susceptible of a fine polish. The tree grows rapidly in England and forms a very ornamental one. The bark of both Brucea antidysenterica and Samatrana is intensely bitter. The former is considered in Abyssinia a sovereign remedy in dysentery and diarrhoea, and Roxburgh was in hopes that the Malayan one would, from the similarity of its sensible properties, be found equally useful in the cure of these complaints. The salt called Brucine, erroneously supposed to be derived from B. antidysenterica, a most acrid and powerful poison is now ascertained to have been obtained from the bark of Nux vomica or Strychnos, all the statements therefore that have been made concerning the danger of using Brucea and Brucine, belong to Strychnos, and have nothing to do with Brucea itself. — Lindley.

Remarks on Genera and Species. In our Prodromus we expressed doubts as to whether Ailanthus malabaricus was really distinct from A. excelsa. Specimens of leaves and flowers of the latter tree, commuaicated by the late Mr. Graham, of Bombay, corresponding with Rheede's figure enable me to testify to that extent, the correctness of his representation. The mature fruit I have not seen but the ovary differs somewhat from the generic character. Ifc has three compressed distinct very obtuse carpels free from the base to the origin of the styles that is about two-thirds of their length : the styles are united throughout from the base to the apex and end in a large 3-lobed spreading stigma. It hence appears, that since the ovaries are distinct that the mature Samara can scarcely be united. In the earliest stages however, they are linear and rounded at the apex as in the plate. The specific character may be improved by the insertion of the words " ovaries 3, distinct, to the origin of the styles, rounded at the apex ; style 1, short, thick, triangular, terminating in a broad spreading 3-lobed stigma, leaflets glabrous on both sides, glaucous beneath."

EXPLANATION OF PLATE 67.

1. Ailanthus excelsa, male — flowering branch, nearly 5. The same magnified but not well represented. natural size. 6. Cluster of carpels — natural size.

2. A male flower seen from above. 7. A carpel cut transversely, showing the solitary

3. Anthers back and front views. seed.

4. An ovary — natural she.

L.—SIMARUBEAE.

This is a small tropical order of which two or three species only have been found on the continent of India; the remainder being from America, continental Africa, and Madagascar.

They are most readily distinguished from the preceding orders, by the insertion of their stamens which in Simarubeae arise from the back of hypogynoas scales like those of Zygophylleae, from which however they are distinguished by their distinct, not. united, ovaries, and from Xanthoxylaceae by the absence of albumen in their seed. They are principally trees.

" Flowers usually bisexual, sometimes unisexual, regular. Calyx 4-5, divided. Petals as many as the sepals and alternate with them, hypogynous, spreading or connivent into a kind of tube : aestivation twisted. Stamens twice as many as the petals : each filament arising from the back of a hypogynous scale : anthers opening longitudinally. Torus a gynophore, bearing the stamens round its base, and the varies on its summit. Ovaries 5 : ovule solitary in each cell, suspended from the inner angle near the apex : styles distinct at the base, but above it uniting into one : stigmas 4-5, distinct, or combined. Fruit of 4-5 indehiscent drupes, arranged round the summit of the gynophore. Seeds pendulous : testa membranaceous. Albumen none. Radicle superior, short, partly concealed within *he thick cotyledons. — Shrubs or trees. Leaves alternate, exstipulate, without dots, simple or compound."

Affinities. They are allied to Rutaceae and its suborders as well as to Connarnceae and Ochnaceae, by their distinct ovaries and gynobase, but are distinguished from the former by the insertions of their stamens, and from the latter by their pendulous, not erect, ovules, and anthers, not opening by terminal pores, to which however they approach through their exalbuminous seed — Adr. de Jussieu remarks, they are known from all Rutaceous plants by the co existence of these characters ; namely, ovaries with but one ovule, indehiscent drupes, exalbuminous seeds, a membranous integument of the embryo, and by the radicle being retracted within thick cotyledons.

Geographical Distribution. The similarity of the Floras of the western coasts of the Indian peninsula of Ceylon and of the Malayan peninsula, is in many cases very remarkable, and in no case more so than in this : the only two Indian species of the order being found in these localities. Samadew indica is a native of Malabar and Ceylon, while S. lucida is from Martaban. There is one, most probably the latter, cultivated in the gardens about Batavia in Java. A third plant doubtfully referred to this order is found on the Himalayas, but forms a distinct genus.

Properties and Uses. Of this order the well known Quassia so remarkable for its pure and intense bitter is a member, and in that property all the other species partake in a greater or less degree. One species a native of Brazil, is said to be so intensely bitter that no insects will attack it. The Himalayan plant referred to possesses the peculiar bitterness of the family in an eminent degree, as does both the seed and the bark of the plant here figured.

EXPLANATION OF PLATE 68.

1. Samadera indica, flowering branch— natural she.

2. A flower partially dissected to show the insertions of the stamens and the ovary.

3. A detached stamen with its attached scale.

4. Stamens and petals removed, showing the elevated gynobase and the ovaries.

5. Ovaries cut transversely, showing the gynophore to which they are attached in the centre.

6. An ovary removed and cut vertically, to show the attachment of the ovule.

7. A cluster of full grown fruit.

8. One of them cut transversely.

9. A seed removed from the capsule.

ZANTHOXYLACEÆ

TODDALIA ACULEATA. (PERS.)

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