CYATHEA—CYCADALES phuric acid, there being no decomposition unless the action is prolonged. When heated it becomes converted into cyanic acid, this reaction being one of the tests that are employed for its detection. CYATHEA, a genus of arborescent ferns, family Cyatheaceae, characterized by having the spores, which are borne on the back of the frond, enclosed in a cup-shaped indusium. There are many species scattered over the tropical regions of the world. C. medullaris is a fine New Zealand species of comparatively hardy character. The soft, pulpy, medullary substance in the centre of the trunk is an article of food somewhat resembling sago. This species and C. dealbata, from the same country, are cultivated as ornamental plants. C. arborea is a West Indian species. CYATHOMETER, an apparatus for determining the level and volume of liquids in closed vessels. It is adapted to bottles and to stationary or movable vessels and is designed especially to prevent fraud in the retail trade in valuable liquids. CYBELE, sib'ě-lē, a goddess of Asia Minor, like Isis, the symbol of the moon, and what is nearly connected with this, of the fruitfulness of the earth, for which reason she is confounded with Rhea, whose worship originated in Crete, and in whom personified nature was revered. According to Diodorus, Cybele was the daughter of the Phrygian king Mæon and his wife, Dindyma. At her birth her father, vexed that the child was not a boy, exposed her upon Mount Cybelus, where she was nursed by lions and panthers and afterward found and brought up by the wives of the herdsmen. She invented fifes and drums, with which she cured the diseases of beasts and children, became intimate with Marsyas and fell violently in love with Attis. She was afterward recognized and received by her parents. Her father discovering her love for Attis had him seized and executed, and left his body unburied. The grief of Cybele on this occasion deranged her understanding and she began a long search for Attis. In art her original statue was nothing but a dark quadrangular stone. Afterward she was represented as a matron, with a mural crown on her head in reference to the improved condition of men arising from agriculture and their union into cities. A common attribute of the goddess is the veil about her head, which refers to the mysterious and incomprehensible in nature. In her right hand she often holds a staff, as an emblem of her power, and in her left a Phrygian drum. Sometimes a few ears of corn stand near her. The sun also is sometimes represented in her right hand, and the crescent of the moon in her left. We sometimes see her in a chariot drawn by lions; or else she sits upon a lion, and, as omnipotent nature, she holds a thunderbolt; or a lion lies near her. These symbols are all representations of her dominion and of the introduction of civilization by her means in the period of barbarism. Her cult was centralized at Phrygia, whence it found its way into Greece, as early as the latter half of the 6th century B.C., and was introduced at Rome in 204 B.C. To the Romans she was known as the Great Idean Mother of the Gods. Under the Roman em 351 pire, it became one of the three most important Roman cults and was of the last pagan worships to give way before Christianity. Consult Farnell, Cults of the Greek States' (Vol. III, Oxford 1907). CYCADACEÆ. See CYCADALES. CYCADALES, sik-a-dā'lēz (from Cycas Neo Lat. nom. pl. of Gr. Kúкaç; the original name of the African cocoa-palm), a greater group of tropic to sub-tropic naked-seeded or gymnospermous plants, the cycads. The large pith of the thick palm-like trunk of certain species of the genus Cycas is the source of the sago starch of commerce, whence the common name, "sago palm." The existing forms are only an ornate remnant of an ancient and varied alliance, the nearest living relative being the "Ginkgo," or maidenhair tree. At least two distinct Cycadalean types are recognizable; the cycads proper include all extant forms and have only a short fossil record, while the cycadeoids, now wholly extinct, were cosmopolitan throughout all of Mesozoic time. The first group is comprised within the order Cycadaceæ, to which all cycadaceous types were until recent years supposed to belong. The second group may be arbitrarily brought within the order Bennettitacea, with far the broader relationships and an immensely varied history. But it appears that variants of both groups go back to the Carboniferous. I. Cycadaceæ.-The cycads are a primitive megaphyllous and composite type with wood structure like conifers (and Cordaites), certain frond and other characters of ferns, and the outward habit of palms. As in the latter, the stem elongates by the slow growth of a terminal bud, with the unfolding of successive crowns of leaves or fronds spirally arranged in close order. As the leaves wilt down, there is formed from their bases an outer more or less persistent armor, which gives the stem its very characteristic appearance. For trunk-forming plants, the cycads are mostly small or even pygmic. They include tuberous to columnar sparsebranched forms, and vary in size from underground trunks a few centimeters in diameter, with fronds no more than a decimeter long, to moderately tall forest forms. It was once suggested that Cycas (see Fig. 2), in which the so-called carpellary leaves alternate with the foliage leaves in armor formation, grew taller than forms bearing cones only. But such is not the fact. The Australian Macrozamia Hopei reaches a height of 60 feet, or nearly double that of any Cycas, and the Mexican Dion spinulosum is nearly as tall. Both these are columnar, rather than branched types. The most robust trunks may reach a meter in diameter, and the longest fronds a length of four meters. Several Central American and Peruvian species are epiphytic. Most of the cycads branch one or more times after reaching a certain age, and all are handsome and easily grown greenhouse plants. The Japanese gardeners display examples of their native species Cycas revoluta, upward of 300 years old, and as freely branched as screw pines. A pot-type is shown in Fig. 3. One of the most interesting of all species is the Cuban Microcycas calocoma, a slender stemmed forest form about 30 feet high, also branched in a manner recalling the habitus of the screw pines. The FIG. 3.- Branching in Cycads. (a), Cycas revoluta, pot-trained in Japan; The fern-like and subterranean stemmed Stan- FIG. 4. Encephalartos, growing in rocky clefts, Lado near Uganda border.- Globular branched trunk type recalling the cycadeoids. Note the two large seed cones. bium and bast, enclosing a large medulla and enveloped by a thick cortex, this being in the main the arrangement in all the gymnosperms and the dicotyls. But there is this difference; bark. Moreover, in Cycas, Macrozama and Encephalartos the woody cylinder does not as in the other genera remain single. After a time the primary cambium becomes inactive, and there successively arise in the cortex secondary cambiums of diminishing power and regularity. From these are produced the so-called "anomalous wood zones," rarely increasing to a dozen and finding no parallel amongst modern plants. The principal features of this second or polyxylic trunk type are shown in Fig. 2, together with a medullar system of anastomosing cauline bundles. These also occur in the pith of Encephalartos. Both pith and cortex are traversed by anastomosing mucilage canals. The cortical bundle system is a complex one, varying greatly in the several genera, and including "girdle leaf traces" (cf. g, Fig. 5) of a primitive, partly concentric structure. The leaf traces are always double and the two branches may nearly girdle the stem. They are most nearly direct in their course from the woody cylinder through the cortex to the leaf base in Zamia and Stangeria. Complexity of the leaf traces begins even in the cotyledonary plate, but is partly a result of the thickness of cortex. The cycads, unlike all vascular cryptogams, send down a primary root which continues as a tap-root. In the case of such subterranean trunks as those of certain species of Zamia (Fig. 1), the tap-root remains prominent, and its lateral branches are relatively small, the trunk assuming a carrot-like form. But in most genera the root system comes to be quite filamentous, being largely made up of freely branching adventitious roots. In Cycas the rootlets come to the surface as a felt-like mass and bear numerous coraloid tubercles, recalling the nitrogen fixing tubercles of legumes. The leaves or fronds are usually of two kinds, scale Fig. 1. Zamia floridana, Miami, Florida, middle of November. (1, to right) ovulate plant with enlarged view of cone, section of the same showing seeds, megasporophyll bearing two seeds, and pinnule; (2) summit of staminate plant bearing five cones; (3) microsporophyll with enlarged view of triple group forming sorus. The entire plant is shown about one-tenth natural size. The staminate plant is about one-third Fig. 2. Cycas revoluta, both columnar and branched stems. Ryûgeji Temple garden, near Fjiri, Japan 1 A Petrified Cycadeoidea from the Black Hills, weight over 300 pounds 2 Restoration of Climbing or Procumbent, Flowering Wielandiella from the Rhaet of Skone, by Nathorst. figures are about one-sixth natural size Both CYCADALES and foliage. There are also the fertile or carpellary leaves of Cycas. All appear in terminal rosettes in the order named, scale, foliage, and when present, carpellary leaves. The scale leaves are dry and aborted foliage leaves, and are present in all but certain species of Macrozamia. The elliptical to acuminate foliage leaves are bipinnate in Bowenia, and pinnate in all the other genera. Prefoliation is direct in Dion and Macrozamia. In Cycas the rachis is straight, but the pinnules are circinnately rolled like those of ferns. Conversely in several other genera the pinnules lie straight along the once deflected rachis. The pinnules vary from broad forms of papery thinness in some Zamias to leathery and hard spinose types like Encephalartos. They vary much in size and form and are of dichotomous venation, except in Cycas, which has a single mid-vein. Those of Stangeria are very fern-like, and Cycas Micholitzii is bilobately dichotomous, signally recalling the fossil Ginkgoid leaves, especially of the Rhätic. The anatomical structure is much as in certain conifers and the extinct Cordaites. Fructification.- The living cycads are all diæcious. The microsporophylls are always organized into cones which may vary from a few centimeters long to enormous forms exceeding in size the very largest ears of Indian corn, and bearing as many as 600 microsporophylls, these being the most massive to be seen in seed plants. In number the staminate cones FIG. 5.- Macrozamia Fraseri. Transverse section of trunk X. (From Worsdell). P. periderm in successively formed layers; w, wall of one of the leaf bases; , leaf base; c, cortex; g, girdle leaf traces; a", isolated bundle of a 3d anomalous wood zone; a', 2d anomalous wood zone; a, 1st anomalous wood zone; n, normal wood (xylem dark, phloem light); m, medulla; m', medullary bundle. vary from a few, or even a single subterminal cone, to over 100 laterally borne cones in the VOL. 8-23 353 Australian Macrozamia Moorei. The sporangia are borne on the under side of the sporophyll and are structurally much like those of Angiopteris among the Marattiacea. In certain genera there is an obscure grouping into sori. (See Fig. 1). The ovules, which are the largest known in the vegetable kingdom, are in the genus Cycas, doubtless the most primitive type among recent phanerogams, borne on the margins of modified leaves, emergent in regular series like the ordinary foliage leaves. In the other genera the megasporophylls each bear but a single pair of ovules, and are organized into terminal, subterminal or lateral cones. These are of striking size in all the genera, and reach a length of nearly a meter with a weight of 90 pounds in some Australian Macrozamias. In various forms following development of a single cone, the stem is prolonged from a new lateral bud as a sympode. But cones truly lateral occur in both Macrozamia and Bowenia, often in some number. The seeds are often of large size. In Cycas, especially, the free seeds look much like large plums; but in cone-bearing forms there are strong appression faces. The seed coat is amphivascular- that is, with a midstone separating an inner and outer bundle system. Cycas is platyspermic, and the other genera have radio-symmetric seeds. There are from one to three cotyledons. Testal structures are closely analogous to those of Ginkgo and the Cordaitales. Even stronger is the resemblance to the seeds of the Paleozoic quasi-ferns, or Cycadofilices, if, as seems necessary, the apical testal division seen in these ancient seeds is regarded as primitive. The seeds of cycads, like those of Ginkgo, are edible. They even form for a considerable part of the year the main food of some of the native Australiansalthough it is said that as freshly gathered the seeds contain a poisonous principle. This is removed by maceration and drying. Fecundation. Among the various primitive characters of the cycads going to prove their descent from homosporous tree-ferns, easily the most recondite is the occurrence of motile multiciliate male cells of the coiled type, characteristic of all the Pteridophytes except the club mosses. These spermatozoids were doubtless common to all the cycadales and are present in Ginkgo, but are not known in any other phanerogams. The pollen grains are first drawn through the micropylar tube into the pollen chamber (by suction), after which the pollen tube ruptures the exine and enters the nucellar tissue, where it may branch. Meanwhile the spermatozoids (Figs. 6, 7) form from the generative cells and, after the rupture of the pollen tube, swim actively to the archegonium through a liquid medium, afforded in part by the tube and probably also by extrusion from the eggcell. The mature spermatozoids are the largest known in any plant (or animal), at least in Zamia floridana. In this species they are visible to the naked eye, and have been studied alive in sugar solutions. They are of nearly spherical top-shape, with a ciliferous spiral running from the apex to the middle region, and motion is mainly by means of their cilia, but there is also amoeboid motion of the spiral end (see Figs. 6 and 7). From this most primitive form of fecundation known in flowering plants it seems evident that not until the later stages of plant |