and Gymnosperms belonging almost entirely to families now extinct, were abundant, while the Pteridophyta attained a development exceeding anything that they can now show. Among the lower classes of plants we have scarcely any knowledge of Palaeozoic Bryophyta; Fungi were probably abundant, but their remains give us little information; while, even among the Algae, which are better represented, well characterized specimens are scanty.
With few exceptions, the remains of Palaeozoic Algae are of comparatively little botanical interest. A vast number of “species” Algae. have been described, but, as has been said, “by far the greater number of the supposed fossil Algae have no claim to be regarded as authentic records of this class of Thallophytes” (Seward, 1898). The investigations of Nathorst, Williamson and others have shown that a very large proportion of the casts and impressions attributed to Algae had in all probability a totally different origin. Some represent the tracks or burrows of worms, crustaceans or other animals; others, the course of rills of water on a sandy or muddy shore; others, again, the marks left on the bottom by bodies drifted along by the waves. In cases of doubt, evidence may be obtained from traces of organic structure, from the presence of carbonaceous matter, or, as Zeiller has pointed out, by the remains of animals such as Bryozoa being attached to the cast, showing that it represents a solid body and not a mere cavity or furrow. Evidence from traces of organization is alone conclusive; the presence of carbonaceous matter, though a useful indication, may be deceptive, for the organic substance may have been derived from other sources than the body which left the impression. The mere external form of the supposed Algae is rarely so characteristic as to afford satisfactory evidence of their nature. Some of the better-attested examples, among which are a few of considerable interest, may now be considered. Of Cyanophyceae, as we should expect, the Palaeozoic remains are very doubtful. Gloioconis, found by Renault in a coprolite of Permian age, was regarded by him as a Cyanophycean allied to Gloeocapsa; this may be so, but the argument drawn from the absence of nuclei, considering the extreme rarity of recognizable nuclei even in the best preserved fossil tissues, can hardly be taken seriously. Girvanella, found in Cambrian, Ordovician and Silurian rocks, as well as in later deposits, appears to have played a part in the origination of oolitic rock-structure. It consists of minute interwoven tubular filaments, and has been variously interpreted as possibly representing the sheaths of a Cyanophycean Alga, and as constituting a Siphoneous thallus of the type of the Codieae. The non-cellular order Siphoneae is fairly well represented in Palaeozoic strata, especially by calcareous verticillate forms referable to the family Dasycladeae; the separate tubular joints of the articulated thallus, bearing the prints of the whorled branches, are sometimes cylindrical (Arthroporella, Vermiporella, &c.), sometimes oval (Sycidium) or spherical (Cyclocrinus). These forms, and others like them, go back to the Silurian and Ordovician; while Gyroporella, from the Permian, is another fairly characteristic Siphoneous type. There can be no doubt that the verticillate Siphoneae, a group much isolated among recent organisms, are among the most ancient families of plants. The gigantic Nematophycus, to be described below, has been regarded as having Siphoneous affinities. Little trace of Confervaceae has been found; Confervites chantransioides, apparently consisting of branched cellular filaments, may perhaps represent a Cambrian Confervoid. Cladiscothallus, from the Culm of Russia, in which the filaments are united to form hemispherical or globular tufts, has been compared by Renault to a Chaetophora. This is one of the somewhat doubtful Algae occurring in boghead coal or torbanite, a carbonaceous rock the nature of which has been much disputed, in the law courts as well as in scientific literature. The boghead of Scotland, Autun and New South Wales is regarded by Renault and Bertrand as mainly composed of gelatinous Algae (Pila and Reinschia), having a hollow, saccate thallus formed of a single layer of cells. It may appear surprising that a body containing 65% of carbon should be so largely made up of gelatinous Algae in a comparatively little altered condition, but the material is rich in bitumen, which seems to have replaced the water contained in the organisms when alive. It has recently been stated, however, that the supposed Algae are in reality the megaspores of Vascular Cryptogams. Scarcely anything is known of Palaeozoic Florideae; Solenopora, ranging from the Ordovician to the Jurassic, resembles, in the structure of its thallus, with definite zones of growth. Corallinaceae such as Lithothamnion, and may probably be of the same nature. A branched filamentous organism from the Lower Carboniferous of Scotland, described by Kidston under the name of Bythotrephis worstoniensis, shows some remains of cellular structure, and may probably be a true Alga, resembling some of the filamentous Florideae in habit.
Apart from the multitude of supposed fossil Algae described as “Fucoids” but usually not of Algal nature, and never presenting determinable characters, very little remains that can be referred to Palaeozoic Brown Algae. The most striking of all fossil Algae, however, Nematophycus, may possibly be a Phaeophycean. The first species of the genus, Nematophycus Logani, was discovered by Dawson in 1856 in the Lower and Middle Devonian of Canada, and was described by him as a Conifer under the name of Protolaxites. Carruthers, however, in 1872 established its Algal nature, and gave it the more appropriate name of Nematophycus. In N. Logani the stem, which is found in a silicified state, may be as much as 3 ft. in diameter. The tissue is made up of large, unseptate, occasionally branching tubes, with an undulating vertical course, among which much smaller tubes are irregularly interwoven. Radially placed gaps in the tissue (at first erroneously interpreted as medullary rays, but subsequently more aptly compared to the air-spaces of large Algae) contain very sparse hyphae, which here branch more freely than elsewhere. The concentric rings of growth, which form a characteristic feature, are due to periodic variations in the size of the larger tubes. Transverse septa have occasionally, but rarely, been detected in the smaller hyphae. Penhallow maintains that these smaller tubes arise as branches from the larger, but other observers have failed to confirm this. In N. Storriei, from the Silurian (Wenlock) of South Wales, described by Barber, there is no sharp differentiation of the two kinds of tubes; they are rarely observed to branch, except in the gaps, which in this species are not radially directed. In N. Ortoni (Penhallow), from the Devonian of Canada, the tubes are quite uniform, and there are no spaces or concentric rings. The tubes have their cavity dilated at intervals, and Penhallow has therefore compared them with the trumpet-hyphae of Laminariaceae, but no transverse septa are anywhere visible. Several other species have been described. Carruthers compared the usually non-cellular structure of Nematophycus with that of Siphoneae such as Halimeda, while recognizing the points of resemblance to Laminariaceae (e.g. Lessonia) in the dimensions of the stem and its concentric rings of growth. Later writers, influenced by the occasional occurrence of transverse walls in the smaller hyphae, have laid more stress on Laminariaceous affinities. The existence of these gigantic Algae in Palaeozoic times, attested by such well-preserved specimens, is a fact of great interest, though their systematic position is still an open question. Pachytheca, a spherical organism, usually about the size of a small pea, found in rocks of Silurian and Devonian age, has been much investigated and discussed, without any decisive light having been thrown on its nature. It was once regarded as connected with Nematophycus (with which it sometimes occurs in association), possibly as its fructification. For this view however, there is no evidence, though the tissues of the two fossils are somewhat similar. Pachytheca is formed of cellular filaments resembling those of a Cladophora, irregularly interwoven in the central region, radiating towards the periphery, and often forked. In one case the spherical thallus was found seated in a cup-like receptacle. There can be little doubt of the Algal nature of the fossil, but beyond this it is impossible at present to carry its determination.
On the whole, it cannot be said that the Palaeozoic remains have as yet thrown much light on the evolution of the Algae, though we may not be prepared to maintain, with Zeiller, that plants of this class appear never to have assumed a form very different from that which they present at the present day.
The first evidence for the existence of Palaeozoic Bacteria was obtained in 1879 by Van Tieghem, who found, that in silicified Bacteria. vegetable remains from the Coal Measures of St Étienne the cellulose membranes showed traces of subjection to butyric fermentation, such as is produced at the present day by Bacillus Amylobacter; he also claimed to have detected the organism itself. Since that time a number of fossil Bacteria, mainly from Palaeozoic strata, have been described by Renault, occurring in all kinds of fossilized vegetable and animal debris. The supposed Micrococci present little that is characteristic; the more definite, rod-like form of the Bacilli offers a better means of recognition, though far from an infallible one; in a few cases dark granules, suggestive of endospores, have been found within the rods. On the whole, the occurrence of Bacteria in Palaeozoic times—so probable a priori—may be taken as established, though the attempt to discriminate species among them is probably futile.
Fungi were no doubt abundant among Palaeozoic vegetation. In examining the tissues of fossil plants of that epoch nothing is Fungi. more common than to meet with mycelial hyphae in and among the cells; in many cases the hyphae are septate, showing that the higher Fungi (Mycomycetes), as distinguished from the more algoid Phycomycetes, already existed. An endophytic Fungus referred to the latter group (Peronosporites antiquarius, W. Smith) bears very definite terminal, or intercalary, spherical vesicles, which may probably be regarded as reproductive organs—either oögonia or sporangia. A minute Fungus bearing sporangia, found by Renault in the wood of a Lepidodendron, and named by him Oöchytrium Lepidodendri, is referred with much probability to the Chytridineae. Conceptacles containing Spores, and strongly suggesting the Chytridineous Fungus Urophiyetis, have recently been found, in petrified material, on the leaves of an Alethopteris, which appears to have undergone decay before fossilization set in. Small spores, almost certainly those of Fungi, are very common in the petrified tissues of Palaeozoic plants. Spherical sacs, bearing forked spines, described by Williamson under the name of Zygosporites, are frequent, usually in an isolated state.
