VII. CHAROPHYTA.
CHARACEÆ. NITELLEÆ.
It has been the general custom to include the Characeæ or Stoneworts among the Chlorophyceae (green algae), of which they form a distinctly isolated family. On the whole, it would seem better to follow the course lately adopted by Migula[437] and allow the Characeæ to rank as a family of a distinct group, Charophyta. While agreeing in many respects with plants higher in the scale than Thallophytes, the Stoneworts do not sufficiently resemble the Bryophyta to be included in that group.
The Charophyta are plants containing chlorophyll, living in fresh and brackish water; the stem is jointed, and bears at the nodes whorls of leaves, on which are borne the reproductive organs. The antheridia are spherical in shape and of complex structure, containing numerous biciliate antherozoids. The oogonia are oval in form and contain a single large egg-cell. The Chara-plant is developed from a protonema formed from the germinating oospore. Vegetative reproduction is effected by means of bulbils, accessory shoots, etc.
The Nitelleæ have not been recognised in a fossil condition. The absence or feeble development of a calcareous incrustation renders the genera of this family less likely to be preserved than such a genus as Chara.
Chareae.
Leaves and stems with or without a cortical investment. Fruit with a five-celled corona. The envelope of the ‘fruit’ and other parts of the plant are frequently encrusted with carbonate of lime.
In the genus Chara, the best known member of the family, the plant as a whole resembles in its general habit and external differentiation of parts the higher plants. The stem consists of long internodes separated by short nodes bearing whorls of leaves. Each internode consists of a long cylindrical cell, which becomes enclosed by a cortical sheath composed of rows of cells which have grown upwards and downwards from the peripheral nodal cells. The cortical cells are usually spirally twisted and impart to the stem a characteristic appearance; they are divided by transverse walls into numerous cells some of which occasionally grow out into short processes (fig. 45 c). The leaves repeat on a smaller scale the structural features of the stem, but possess a limited growth, whereas the stem has an unlimited power of growth by means of a large hemispherical apical cell. Branches arise in the axils of the leaves. The plants are either monoecious or dioecious. The oogonium is elliptical in shape, and is borne on a short stalk-cell, it contains a single oosphere. The wall of the oogonium is formed of five spirally twisted cells which have grown over it from the five peripheral cells of a leaf-node. The tips of the investing cells project at the apex in the form of a terminal crown or corona (fig. 45, E, c). The antheridia have a complex structure, and produce a very large number of motile antherozoids.
Fig. 45. A and B. Chara Knowltoni Sew. From a section in the British Museum. C. Stem of Chara foetida A. Br. in transverse section (after Migula. × 18). D. Interior of oogonium of C. foetida. E. Oogonium of C. foetida (D and E after Migula. × 50).
After fertilisation, the egg-cell becomes surrounded by a membrane, at first colourless, but afterwards yellow or brown. The inner cell-walls of the cells surrounding the oospore become thicker and darker in colour; the outer walls remain thin and eventually fall away. The lateral walls may or may not become thickened. In most of the Chareae a calcareous deposit is formed between the hard shell and the outer walls of the cells enveloping the oospore. This calcareous shell is developed subsequently to the thickening and hardening of the inner walls of the fruit-case. The cells of the corona and stalk do not become calcareous. In the fossil Charas, it is this calcareous shell that is preserved. In the members of the Chareae the stems are usually encrusted with carbonate of lime, and thus have a much better chance of preservation than the slightly calcareous Nitelleæ.
Chara.
The generic characters have already been described in the brief account of the family Chareae.
The generic name was proposed by Vaillant in 1719[438], and adopted by Linnaeus, who classed the Stoneworts with aquatic phanerogams. As long ago as 1623[439] a figure of Chara was published by Caspar Bauhin as a form of Equisetum. The generic name Chara has usually been applied to recent and fossil species alike. The existing species have a wide distribution; Chara foetida, A. Br., a common British form, occurs in practically all parts of the world. Stems and calcareous ‘fruit-cases’ occur fairly commonly in a fossil state, and differ but little from recent species, at least as regards essential features.
It is difficult to say at what geological horizon the Stoneworts are first represented. The first certain traces of Chara occur in Jurassic rocks, but certain spirally marked subspherical bodies have been recorded from Devonian and Carboniferous strata, which closely resemble Chara oogonia, and may be Palaeozoic representatives of the genus.
In 1889 Mr Knowlton[440] of the American Geological Survey described some ‘problematic organisms’ found in Devonian rocks at the falls of the Ohio. Examples of these fossils are shown in fig. 46 b and c; the spirally grooved body measures from 1·50 to 1·80 mm. in diameter, and about 1·70 mm. in length. The Chara-like character of the fossils had been previously suggested by Meek[441] in 1873. Without going into the arguments for or against placing these fossils in the Chareae, they may at least be mentioned as possible but not certain Palaeozoic forms of Chara or an allied genus.
Fig. 46. a. Chara Bleicheri Sap. × 30. b and c. Devonian Chara? sp. circa × 12. d and e. Chara Wrighti Forbes. circa × 12.
1. Chara Bleicheri, Saporta. Fig. 46, a.
In this form the ‘fruits’ are minute and subspherical, ·39-·44 mm. long, and ·35-·40 mm. broad, showing in side view 5–6 slightly oblique spiral bands. Each spiral band bears a row of slightly projecting tubercles.
This species was first described by Saporta[442] from the Oxfordian (Jurassic) rocks of the Department of Lot in France; it is compared by the author of the species with Chara Jaccardi Heer, described by Heer from the Upper Jurassic rocks of Switzerland.
2. C. Knowltoni, Seward. Fig. 45, a and b, and Fig. 47.
The Oogonia are broadly oval, about ·5mm. in length, and at the broadest part of about the same breadth. The surface is marked by eleven or twelve bands in the form of a flattened spiral. The stems possess investing cortical cells.
This species was founded on specimens from the Wealden beds of Sussex[443], but numerous examples of Chara ‘fruits’ and stems have long been known from the uppermost Jurassic rocks of the Dorset coast and the Isle of Wight, which may probably be included in this species. These fossil Charas are abundant[444] in the Chert beds of Purbeck age seen in the cliffs near Swanage. Pieces of corticated stems from this locality are represented in fig. 45 A and B.
The cortical cells surrounding a large internodal cell are very clearly seen in the section shown in fig. 45 B, and in the longitudinal view in fig. 45 A. The resemblance of these specimens to the stems of recent Stoneworts is very striking.
Fig. 47. Chara Knowltoni, Sew. × 30.
The single oogonium of fig. 47 was found in the Wealden beds near Hastings.
3. Chara Wrighti, Forbes. Fig. 46, d and e.
This species is characterised by globular or somewhat elliptical oogonia, with six or seven spiral bands.
It is very abundant in the Lower Headon beds of Hordwell Cliffs on the Hampshire coast[445]. Various species of Chara are commonly met with in the Oligocene beds of the Isle of Wight and Hampshire, as well as in the Paris basin beds, and elsewhere. Well preserved ‘fruits’ and stem fragments are met with in a siliceous rock of Upper Oligocene age imported from Montmorency in the Paris basin, and used as a stone for grinding phosphates at some chemical works near Upware, a few miles from Cambridge.
Many other species of fossil Charas are known from various horizons and localities, but the above examples suffice as illustrative types. In Post-Tertiary deposits masses of Chara and plant fragments occasionally occur forming blocks of Travertine. Examples of such Chara beds have been recorded by Sharpe from Northampton[446], by Lyell[447] from Forfarshire, and by other writers from several other districts. Beds of calcareous marl are occasionally seen as whitish streaks in the peat of the Fenland[448]; these often consist in great part of Charas. A season’s growth of Chara in a shallow lake or mere in the Fens may appear as a white line in a section of peaty and other material which has been formed on the site of old pools or lakes.
The recognition of specific characters in the isolated Chara ‘fruits’ usually met with in a fossil state is exceedingly unsatisfactory; the features usually relied on in the living species are not preserved, and great care should be taken in the separation of the various forms.