ORDER XXVI.-BALANOPHORACEÆ.-CYNOMORIUMS.

Balanaphores, Rich. in Mem. Mus. 8, 429. (1822); Endlicher Meletemata, p. 10. (1832); gen. xxxix. Meisner, p. 366; Junghuns in nov. act. xviii. suppl.; Griffith, Proceedings Linn. Soc. No. xxii. DIAGNOSIS.-Stems amorphous, fungoid; peduncles scaly flowers in spikes; ovules solitary, pendulous; fruit one-seeded.

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Fig. LXIII.

Leafless, brown, red, white, or yellow (never green) root-parasites, with underground fleshy horizontal branched rhizomes, or more generally amorphous tubers, from which spring erect simple (rarely branched) peduncles, that are naked, or covered with scattered or imbricating scales, rarely combined into an involucre. Flowers red, yellow, or white, unisexual (rarely bisexual), monoecious or dioecious, collected into dense, spherical or cylindrical, entire, lobed, or branched heads, often mixed with simple articulated filiform or club-shaped filaments. Bracts very variable, or absent; sometimes, when the heads are lobed, large, peltate, and imbricated, each subtending and often covering a lobe (branch) of the head; at other times the bracts are scattered promiscuously amongst the flowers; sometimes they are peltate, and connected by their contiguous edges into an areolate indusium, that falls away piece

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meal as the head enlarges; at others the flowers are arranged on their stipes. Male flowers conspicuous, usually white, pedicelled, exserted beyond the filaments and female flowers, generally at the base of hermaphrodite heads, or scattered irregularly amongst female flowers, rarely wholly naked, then consisting of anthers crowded on a branched spike. Perianth tubular or funnel-shaped, entire or split, or more frequently 3-5-lobed, with valvate aestivation; lobes patent, or reflexed, fleshy, white, or highlycoloured. Stamens usually 3-5, with both filaments and anthers more or less connate or free, the latter frequently forming a lobed 6-12-celled mass, bursting outwards, or rarely inwards. (Stamen solitary, epigynous, and introrse, in bisexual flowers of Cynomorium. Stamens 3, nearly free and extrorse in Langsdorffia; 3 and free in Sarcophyte, where each filament bears a capitate anther, that breaks up into a many-celled mass. Anthers

Fig. LXIV.

Fig. LXIII. Scybalium fungiforme. 1. A male plant; 2. a female; 3. male flowers with hairs between them; 4. females; 5, a vertical section of a female, with the two pendulous ovules; 6. a section across a ripe fruit.

Fig. LXIV. Cynomorium coccineum. 1. A section of the ripe fruit, showing the embryo on the right of the albumen; 2. a portion of the nucleus very highly magnified, showing the embryo and the angular cells among which it lies. N.B. These cells are separated by the pressure of a compressorium.

numerous and anfractuose in Polyplethia.) Pollen globose, yellow, 1-3-nucleate. Female flowers very minute, densely crowded, sessile or stalked, sometimes seated round the base of a club-shaped pedicel (Balanophora), shorter than the filaments amongst which they often nestle, and beyond which the styles protrude; generally consisting of a compressed ovarium, with 1-2 styles. Perianth seldom apparent, tube closely investing the ovary, and not distinguishable from it; limb 2-lipped, or none; rarely (Cynomorium), there are 6 irregularly inserted valvate pieces of the perianth. Ovary 1-celled (2-celled according to Endlicher in Scybalium). Ovule a solitary naked pendulous nucleus; position of apex unknown. Styles 1 or 2, filiform, each with a simple papillose stigma. (Style flattened in Cynomorium, having two parallel chords, and two papillose stigmatic points. In Sarcophyte, the female flowers coalesce into fleshy capitula, which are sessile on a branched axis, and have sessile stigmata.) Fruit a small compressed nut; epicarp rather fleshy; endocarp crustaceous. Seed solitary, pendulous, filling the cavity of the pericarp. Albumen of large, hard, densely packed grains, adherent to the delicate membrane that surrounds them. Embryo lateral in Cynomorium and Corynæa; spherical, undivided, and soft in the former; harder, compressed, and lobed in the latter.

"A very remarkable natural order, displaying much variety in habit and structure of the floral organs, but agreeing in all essential characters, especially in those of reproduction. They have been likened to Fungi in appearance and mode of growth by some, but others fail to recognize any such similarity. They differ wholly from that natural order in consistence, anatomy, structure, slow mode of growth, and in having conspicuous brightly coloured male flowers.

"The earliest stage at which I have examined Balanophora, and some allied American plants, presents a minute amorphous cellular mass, nestling in ruptures of the bark of the root of the plant, which is henceforward the stock. Vascular tissue forms in the axis of this mass, which swells, and displacing more bark, enlarges to a tuberous rhizome, that in most cases finally envelopes the root, to which it is attached on one side only. In Helosis and Langsdorffia, a branched rhizome is formed underground. This rhizome has a dicotyledonous arrangement of its vascular plates, and sometimes gives off rootlets, which, when they come in contact with other rootfibres of the stock, induce a specific action on them, terminating in destruction of the bark, and a further attachment of the parasite. In no case has the germination of the embryo been observed, but the subsequent stages of growth so entirely resemble those of Loranthaceæ, that there appears no reason to expect any anomaly in the first stages.

"The rhizome, when tuberous, is generally covered with large lenticels (?), consisting of cellular, often cruciform pustules, uncovered by the cuticle. In these tubers the arrangement of the vascular bundles into plates is also exogenous, resembling closely in anatomical details that which prevails in some species of Loranthaceæ. Bundles of vascular tissue run from the tuber (or rhizome) into the peduncle, where they are often symmetrically arranged, and supply the scales, the bracteæ, and the lobes or branches of the inflorescence. The cellular tissue is composed of large nucleated cells, full of resinous matter (and some starch grains?); the vascular of woody tubes; large, barred, spirally-marked, seldom unrollable tubes, of bothrenchyma, and of cylindrical and hexagonal, simple and septate tubes; also of copious thick sclerogen tubes and cells, with perforated walls. There are no true spiral vessels, and the vascular tissue is always in contact with the wood of the stock, which latter sometimes ramifies in the tuber. This contact is sometimes so intimate, that it is impossible to separate the vascular bundles of the parasite from those of the stock; and after long maceration, the latter may hence be traced apparently running continuously up the peduncle, and into the head of flowers. The peduncle bursts from any part of the tuber or rhizome; in its youngest state it is generally covered by imbricating scales; vascular bundles form independently in its substance, and descend to join those of the rhizome. The cuticle of the peduncle and scales has no stomates, whose functions are probably performed by the lenticels of the rhizome. The filaments that nestle amongst the female flowers of the American genera, and of Phæocordylis, appear to be abortive female flowers in most cases, but may, in these and others, be also in part reduced, deformed, or displaced segments of a perianth, as analogy with Cynomorium suggests. They have been compared with the paraphyses of mosses,* with which they have no further affinity or analogy than that both are cellular organs. The ovaria have also been likened to the pistillidia of Mosses and Hepatica, to which they bear no relation in structure, origin, or function. In the youngest state, the female flowers of some American species are 2- and even 3-lobed, with as many styles.

* Griffith, Linn. Soc. Trans. v. 20, p. 99 and 102.

"In endeavouring to determine the affinities of Balanophoreæ, I have confined my attention to the organs of reproduction, which, whether male or female, are perfect, and typical of Phænogamic plants in all respects, though reduced in number and proportion of parts; at the same time rejecting the more prominent, but comparatively unimportant characters of growth and appearance, colour, parasitism, and the inability of most observers to find embryos in some, such being conspicuous in others. The exogenous arrangement of the vascular bundles of the rhizomes, lobed embryo of Corynea, and decidedly dicotyledonous one of Mystropetalum are the most important positive characters hitherto observed, by which to determine the division of the vegetable kingdom to which Balanophores should be referred, which is hence Dicotyledones. The one-celled inferior ovarium, with often two styles, adherent two-lipped or truncate perianth, unisexual flowers, epigynous stamen when the flowers are bisexual, solitary pendulous ovule, and the structure of the albumen and embryo wherever these are made out, are all typical of plants referred to or closely related to Halorageæ, an order in which there is a great tendency to imperfection of the floral organs. The female inflorescence of Lepidophyton (n. g. ined.) differs in no respects, except in having larger bracteæ, from that of Gunnera scabra; whilst the dense albumen of all Balanophoreæ, composed of large grains covered with a membranous testa, that adheres to the walls of the pericarp, and the minute embryo, also characterize the seed of all species of Gunnera. The bisexual monandrous flowers of Cynomorium in many respects closely resemble those of Hippuris, and its perianth is more highly developed, though more irregular in insertion. Equally strong and decided points of affinity may be found in the male flowers, both in the development and suppression of their parts, but such details are unsuited to these pages.

'Balanophoreæ are found on the roots of various Dicotyledonous plants (Vines, Maples, Oaks, Araliaceæ, &c. &c.) and abound in the mountains of tropical countries, especially the Andes of Peru and Colombia, the Himalaya (where they ascend to 11,000 feet in Lat. 28° N.) and Khasia Mountains of India. In the old world one (Cynomorium) is found in Malta, N. Africa, the Levant, and Canaries; another on the west coast of Africa; Sarcophyte in South Africa. Eight or ten species inhabit the Indian Continent, and others its islands, the north coast of Australia and Polynesia. As many are found in Mexico, Central and South America, and Jamaica (where Phyllocoryne, n. g. ined., is called Jim Crow's Nose). A few are Brazilian, and Mr. Miers informs us that one grows on the Pampas.

"The direct uses are few. They seem, as far as anything is known of them, to be styptics. The Cynomorium coccineum, or Fungus melitensis of the apothecaries, long had a great reputation in that way; and various kinds of Helosis have had a similar character. Sarcophyte, a Cape plant, is said to have an atrocious odour. Pöppig says, that Ombrophytum, which in Peru springs up suddenly after rain, in the manner of the toadstool, is called Mays del Monte, in consequence of its resemblance to a kind of Maize, and is quite insipid, on which account it is cooked and eaten like Fungi. This, if true, presents a remarkable contrast with the Balanophoras of India, whose spikes are very slowly developed, and decay after ripening their seeds very gradually indeed. Lepidophytum is also eaten in Bolivia. Cups, used throughout Tibet, are turned from knots produced on the roots of maples by the Himalayan species."

ORDER XXVII. CYTINACEA.--CISTUSRAPES.

Cytinere, Adolph. Brongn. in Ann. des. Sc. Nat. 1. 29. (1824); Endlicher Meletemata, p. 13. Gen. xl. Meisner, p. 367. R. Brown in Linn. Trans. xix.-Pistiaceæ, Agardh. Aphor. Bot. p. 240. (1826).Aristolochiæ, & Cytineæ, Link Handb. 1. 368. (1829).-Hydnorea, R. Br. Linn. Trans. 19. (1844). DIAGNOSIS.-Flowers in spikes at the end of a scaly stem, with a 3-6-parted calyx, anthers opening by slits, and innumerable ovules growing over parietal placenta.

Flowers, or, solitary and stemless, or clustered at the top of a stalk covered with imbricated scales, the males uppermost, the females lowermost, in the axil of a bract, and supported on each side by a bractlet. Perianth tubular-campanulate, with a spreading 3-6-lobed limb, the segments imbricated, the exterior alternating with the bractlets

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or induplicate and valvate. Anthers sessile, 2-celled; their cells distinct, opening longitudinally; four dissepiment-like membranes in Cytinus alternate with the segments of the perianth, and join its tube with the column. ? Perianth as in the males, but epigynous. Ovary inferior, 1-celled, with vertical or parietal placenta, covered by innumerable ovules; style cylindrical, joined to the tube of the perianth by septiform processes, with a thick stigma, or free, and consisting of several styles, each having a free stigmatic apex. Fruit berried, leathery, one-celled, with innumerable seeds buried in pulp, and having a hard leathery skin firmly attached to the nucleus. Seed in Hydnora, with a small undivided embryo in the centre of cartilaginous albumen, and in Cytinus exalbuminous according to Brown.

In these we have a near approach to the common condition of Endogens, both in structure and habit, if we compare Cytinus with some Bromelworts. But the appearance of Hydnora is so peculiar that we know nothing to contrast it with, except some such Fungus as a Geaster, like which it grows half-buried in the soil. Its innumerable seeds distinguish it from Rafflesiads, as well as its caulescent habit and slit anthers.

Fig. LXV.

The history of this extraordinary plant has been fully given by Ferdinand Bauer and Dr. Brown, in the 19th vol. of the Linnean Transactions, from which place the accompanying cuts are taken. The genus is regarded by Brown as the type of a peculiar Order and perhaps with justice. But for reasons elsewhere given, I demur to the formation of all Orders that depend upon a single genus.

Cytinus is parasitical on the roots of Cistus in the South of Europe; the rest are from the Cape of Good Hope, where Hydnora is parasitical on the roots of succulent Euphorbias, and of Cotyledon orbiculatum.

Hydnora Africana (Jackals Kost or Kauimp), smells like decaying roast-beef, or some fungus (Harvey); when roasted it is eaten by the African savages. Cytinus Hypocistis (oxiri Diosc.) contains gallic acid, and according to Pelletier, has the property of precipitating gelatine without containing tannin; its extract is still officinal in the South of Europe, under the name of Succus Hypocistidis; it is blackish, sub-acid, astringent, and is employed in hæmorrhages and dysentery.

Fig. LXV. Cytinus Hypocistis. 1. A flower; 2. a head of anthers; 3. a transverse rection of the

Ovary.

Fig. LXVI.

Fig. LXVI.

Fig. LXVII.

A plant of Hydnora Africana. Fig. LXVII.-A longitudinal section of it. Ferd. Bauer.

Blume suggests that Thismia (his genus Sarcosiphon) may be the type of a new order of Rhizanths. He describes it as a leafless parasite, growing on the roots of trees, resembling in appearance an Angiogastrous Fungus, and smelling like stinking fish. Miers refers this genus to Triurids (see p. 172).