Jungermanniaceae Acrogynae.—The plant consists of leafy shoots, the origin of which can be understood in the light of the foliose forms described above. The great majority of existing liverworts belong to this group, the general plan of construction of which is throughout very similar. In Britain thirty-nine genera with numerous species are found. With few exceptions the stem grows by means of a pyramidal apical cell cutting off three rows of segments. Each segment gives rise to a leaf, but usually the leaves of the ventral row (amphigastria) are smaller and differently shaped from those of the two lateral rows; in a number of genera they are wanting altogether. Sometimes the leaves retain their transverse insertion on the stem, and the two lobes of which they consist are developed equally. More often they come to be obliquely inserted, the anterior edge of each leaf lying under or over the edge of the leaf in front. The two lobes are often unequally developed. In Scapania the upper lobe is the smaller, while in Radula, Poretta and the Lejeuneae this is the case with the lower lobe. The folding of one lobe against another assists in the retention of water. Pitcher-like structures have arisen in different ways in a number of genera, and are especially common in epiphytic forms (Frullania, Lepidolaena, Pleurozia). In some forms the leaves are finely divided, and along with the hair-like paraphyllia form a loose weft around the stem (Trichocolea). The rhizoids spring from the lower surface of the stem, and sometimes from the bases of the leaves. The branches arise below and by the side of the leaves.
Fig. 9.—Cephalozia bicuspidata. Longitudinal section of the summit of a shoot bearing a nearly mature sporogonium, sg, still enclosed in the calyptra; ar′, archegonia which have remained unfertilized; st, stem; b, leaf; p, perianth. (After Hofmeister.)
The sexual organs may occur on the same or on distinct individuals. The antheridia are protected by leaves which are often modified in shape. The archegonia are borne at the apex of the main stem or of a lateral branch. A single archegonium may arise from the apical cell (Lejeunea); more commonly a number of others are formed from the surrounding segments. The leaves below the archegonial group are frequently modified in size and shape, but the chief protection is afforded by a tubular perianth, which corresponds to a coherent whorl of leaves and grows up independently of fertilization. The perianth serves also to enclose and protect the sporogonium during its development. In a number of forms belonging to different groups the end of the stem on which the sporogonium is borne grows downwards so as to form a hollow tubular sac enclosing the sporogonium; in other cases this marsupial sac is formed by the base of the sporogonium boring into the thickened end of the stem. The sac usually penetrates into the soil and bears rhizoids on its outer surface. Kantia, Calypogeia and Saccogyna are British forms, which have their sporogonia protected in this way. The sporogonium is very similar throughout the group (figs. 8, 9). At maturity the seta elongates rapidly, and the wall of the capsule splits more or less completely into four valves, allowing the elaters and spores to escape. In the Jubuloideae, which in other respects form a well-marked group, the seta is short and the elaters extend from the upper part of the capsule to the base; at dehiscence they remain fixed to the valves into which the capsule splits. The germinating spore usually forms a short filament, but in other cases a flat plate of cells growing by a two-sided apical cell is first formed (Radula, Lejeunea). In one or two tropical forms the pro-embryonic stage is prolonged, and leafy shoots only arise in connexion with the sexual organs. In Protocephalozia, which grows on bare earth in South America, this pro-embryo is filamentous, while in Lejeunea Metzgeriopsis, which grows on the leaves of living plants, it is a flat branched thallus closely applied to the substratum. Other cases of the plant being, with the exception of the sexual branches, apparently thalloid, are on the other hand to be explained as due to the reduction of the leaves and flattening of the stem of a shoot (Pteropsiella, Zoopsis).
The Acrogynous Jungermanniaceae fall into a number of natural groups, which cannot, however, be followed out here. They occur in very various situations, on the ground, on rocks and stones, on tree trunks, and, in the damp tropics, on leaves. Usually they form larger or smaller tufts of a green colour, but some forms have a reddish tint.
Fig. 10.—Anthoceros laevis. sp, Sporogonium; c, columella.
From Strasburger's Text-book of Botany.
Anthocerotales.—This small and very natural group includes the three genera Anthoceros, Dendroceros and Notothylas, and stands in
many respects in an isolated position among the Bryophyta. Three species of Anthoceros occur in Britain, growing on the damp soil of fields, ditches, &c. The dark green thallus has an ill-defined midrib, and is composed of parenchymatous cells. In each assimilating cell there is usually a single large chloroplast. The apical region, which has a single initial cell, is protected by mucilage secreted by the mucilage slits, which are small pit-like depressions between superficial cells of the lower surface. Mucilage is also often formed in intercellular spaces within the thallus. Colonies of Nostoc are constantly found living in some of the mucilage slits which then become enlarged. The sexual organs are scattered over the upper surface. The stalked globular antheridia are exceptional in being formed endogenously, and are situated in groups in special intercellular spaces. The superficial layer of cells bounding the cavity does not break down until the antheridia are nearly mature. Occasionally antheridia develop on the surface of shaded portions of the thallus. The necks of the archegonia hardly project above the general surface of the thallus. In structure and development they agree with other Hepaticae, though differences of detail exist. The young sporogonium is protected by a thick calyptra derived from the tissue of the thallus around the archegonium. The sporogonium consists of a large bulbous foot, the superficial cells of which grow out into processes, and a long capsule, which continues to grow for months by the activity of a zone of cells between it and the foot, and may attain the length of an inch and a half. The wall of the capsule is several layers of cells thick, and since the epidermis contains functional stomata and the underlying cells possess chlorophyll it is capable of assimilation. In the centre of the capsule is a strand of narrow elongated cells forming the columella, and between this and the wall spores mixed with elaters are formed from the dome-shaped archesporium, the origin of which has already been described (fig. 4, D). The capsule opens by splitting into two valves from the apex downwards, and the mature spores escape while others are developing in succession below. In Dendroceros, which grows as an epiphyte in the tropics, the thallus has a well-defined midrib and broad wings composed of a single layer of cells. The capsule is similar to that of Anthoceros, but has no stomata, and the elaters have spirally thickened walls. Some species of Anthoceros agree with it in these respects. Notothylas resembles Anthoceros in its thallus, but the sporogonium is much smaller. In some species, although the columella and archesporium arise in the usual way, both give rise to mingled spores and elaters, and no sterile columella is developed.