Fig. 35, A, affords a diagrammatic view of a Diplopora tube, and shews the arrangement of the numerous whorls of canals. In fig. 35, B, a piece of limestone is represented containing several Diploporas cut across transversely and more or less obliquely. In an obliquely transverse section of a tube perforated by horizontal canals the cavities of the canals necessarily appear as holes or discontinuous canals in the substance of the calcareous wall. The manner of occurrence of the specimens points to the abundance of this genus in the Triassic seas, and suggests that the calcareous tubes of Diplopora may have been important factors in the building up of limestone sediments[323]. In many instances no doubt the carbonate of lime of the thallus has been dissolved and recrystallised, and the original form completely obliterated. As in the rocks built up largely of calcareous Florideae (p. 185) which have lost their structure, it is a legitimate inference that some of the limestone rocks which shew no trace of organic structure may have been in part derived from the calcareous incrustation of various algal genera.
In this genus from the Alpine Trias the structure of the calcareous tube is very similar to that in Diplopora, but in Gyroporella the canals form less distinct whorls and are closed externally by a small plate, as seen in figs. 35, C and D.
As Solms-Laubach has pointed out, the branch-systems of Diplopora, Gyroporella and other older genera are much simpler than in the Tertiary genera Dactylopora and others[324].
A species of Gyroporella, G. bellerophontis, has recently been described by Rothpletz[325] from Permian rocks in the Southern Tyrol. The thallus is tubular in form and has a diameter of ·5–1 mm.
The genus Dactylopora was founded by Lamarck[326] on some fossil specimens from the Calcaire Grossier and included among the Zoophytes. D’Orbigny afterwards included it among the Foraminifera, and the structure of the calcareous body has been described by Carpenter[327] and other writers on the Foraminifera. In a specimen of Dactylopora cylindracea Lam. from the Paris basin, for which I am indebted to Munier-Chalmas, the tubular thallus measures 4 mm. in diameter; at the complete end it is closed and bluntly rounded. The wall of the tube is perforated by numerous canals, and contains oval cavities which were no doubt originally occupied by sporangia. The shape of the specimens is similar to that of Diplopora, but the canals and cavities present a characteristic and more complex appearance, when seen in a transverse section of the wall, than in the older genus Diplopora. Gümbel has given a detailed account of this Tertiary genus in his memoir on Die sogenannten Nulliporen[328]; he distinguishes between Dactyloporella and Gyroporella by the existence of cavities in the calcareous wall of the tube in the former genus, and by their absence in the latter. The oval cavities in a Dactyloporella were originally occupied by sporangia; in Diplopora and Gyroporella the sporangia were probably borne externally and on an uncalcified portion of the thallus.
In addition to the few examples of fossil species described above there are numerous others of considerable interest, which illustrate the great wealth of form among the Tertiary and other representatives of the Verticillate Siphoneae.
Reference has already been made to Vermiporella as an example of a Silurian genus. Other genera have been described by Stolley from Silurian boulders in the North-German drift under the names Palaeoporella, Dasyporella and Rhabdoporella[329]; the latter genus is compared with the Triassic Diplopora, and the two preceding with the recent Bornetella.
Schlüter has transferred a supposed Devonian Foraminiferal genus, Coelotrochium[330], to the list of Palaeozoic Siphoneae. Munier-Chalmas regards some of the fossils described by Saporta under the name of Goniolina[331], and classed among the inflorescences of pro-angiospermous plants, as examples of Jurassic Siphoneae. The shape and surface-features of some of the examples of Goniolina suggest a comparison with Echinoid spines, but the resemblance which many of the forms in the Sorbonne collection present to large calcareous Siphoneae is still more striking. A comparison of Saporta’s fig. 5, Pl. xxxiii. and fig. 4, Pl. xxxii. in volume iv. of the Flore Jurassique, with the figures given by Solms-Laubach[332] and Cramer[333] of species of Bornetella brings out a close similarity between Goniolina and recent algae; the chief difference being the greater size of the fossil forms. The possibility of confounding Echinoid spines with calcareous Siphoneae is illustrated by Rothpletz[334], who has expressed the opinion that Gümbel’s Haploporella fasciculata is not an alga but the spine of a sea-urchin.
Among Cretaceous forms, in addition to Goniolina, which passes upwards from Jurassic rocks, Triploporella[335] and other genera have been recorded.
Uteria[336] is an interesting type of Tertiary genera; it occurs in the form of barrel-shaped rings, which are probably the detached segments of a form in which the central axial cell was encrusted with carbonate of lime, but the sporangia and the whorls of branches differed from those of Cymopolia in being without a calcareous investment.
b. Confervoideae.
Without attempting to describe at length the fossil forms referred to this division of the Chlorophyceae, there is one fossil which deserves a passing notice. Brongniart in 1828[337] instituted the generic term Confervites for filamentous fossils resembling recent species of confervoid algae. Numerous fossils have been referred to this genus by different authors, but they are for the most part valueless and need not be further considered. In 1887 Bornemann described some new forms which he referred to this genus from the Cambrian rocks of Sardinia. He describes the red marble of San Pietra, near Masne, as being in places full of the delicate remains of algae having the form of branched filaments, and appearing in sections of the rock as white lines on a dark crystalline matrix. In fig. 35, G, one of these Sardinian specimens is represented. This form is named Confervites Chantransioides[338]; the thallus consists of branched cell-filaments, having a breadth of 6–7µ, and composed of ovate cells. It is possible that this is a fragment of a Cambrian alga, but the figures and descriptions do not afford by any means convincing evidence. From post-Tertiary beds various genera, such as Vaucheria and others, have been recorded, but they possess but little botanical value.
C. INCERTAE SEDIS.
During the last few years much has been written by two French authors, Dr Renault and Prof. Bertrand, on the subject of the so-called Boghead of France, Scotland, and other countries. They hold the view that the formation of the extensive beds of this carbonaceous material was due to the accumulation and preservation of enormous numbers of minute algae which lived in Permo-Carboniferous lakes.
In an article contributed to Science-Progress in 1895 I ventured to express doubts as to the correctness of the conclusions of MM. Renault and Bertrand[339]. Since then Prof. Bertrand has very kindly demonstrated to me many of his microscopic preparations of various Bogheads, and I am indebted to Prof. Bayley Balfour of Edinburgh for an opportunity of examining a series of sections of the Scotch Boghead. The examination of these specimens has convinced me of the difficulties of the problems which many investigators have tried to solve, but it has by no means led me to entirely adopt the views expressed by MM. Bertrand and Renault.
The Boghead or Torbanite of Scotland was rendered famous by a protracted lawsuit tried in Edinburgh from July 29th to August 4th, 1853. A lease had been granted by Mr and Mrs Gillespie, of Torbanehill, in Fifeshire, to Messrs James Russell and Son, coal-masters of Falkirk, of “the whole coal, ironstone, iron-ore, limestone, and fire-clay (but not to comprehend copper, or any other minerals whatsoever, except those specified) with lands of Torbanehill[340].” After the Boghead had been worked for two years the Gillespies challenged the right of Messrs Russell, and argued that the valuable mineral Torbanite was not included among the substances named in the agreement. The defendants maintained that it was a coal, known as gas-, cannel- or parrot-coal. A verdict was given for the defendants. Some of the scientific experts who gave evidence at the trial considered that the Boghead afforded indications of organic structure, while others regarded it as essentially mineral in origin.
The Torbanite or Boghead is a close-grained brown rock, of peculiar toughness and having a subconchoidal fracture. It contains about 65% carbon, with some hydrogen, oxygen, sulphur, and mineral substances. A thin section examined under the microscope presents the appearance of a dark and amorphous matrix, containing numerous oval, spherical and irregularly shaped bright orange-yellow patches. Fig. 36, 1 shows the manner of occurrence of the yellow bodies in a piece of Scotch Boghead, as seen in a slightly magnified horizontal section. Under a higher power the light patches in the figure reveal traces of a faint radial striation, which in some cases suggests the occurrence of a number of oval or polygonal cells.
The Autun Boghead possesses practically the same structure. The yellow bodies are often sufficiently abundant to impart a bright yellow colour to a thin section. If the section is vertical the coloured bodies are seen to be arranged in more or less regular layers parallel to the plane of bedding.
The Kerosene shale of New South Wales agrees closely with the Scotch and French Boghead; it is approximately of the same geological age, and is largely made up of orange or yellow bodies similar to those of the European Boghead, but much more clearly preserved.
The nature and manner of formation of the various forms of coal should be dealt with in a later chapter devoted to the subject of plants as rock-builders, but in view of the recent statements as to the algal nature of these bituminous deposits it may not be out of place to state briefly the main conclusions of the French authors.
MM. Renault and Bertrand regard each of the yellow bodies in the European and Australian Boghead as the thallus of an alga. To the form which is most abundant in the Kerosene shale they have given the generic name of Reinschia, while that in the Scotch and French Boghead is named Pila.
A section of a piece of Kerosene shale at right angles to the bedding appears to be made up of fairly regular layers of flattened elliptical sacs of an orange or yellow colour. Each sac or thallus is about 300µ in length and 150µ broad (fig. 36, 3). A single row of cells constitutes the wall surrounding the central globular cavity. The cells are more or less pyriform in shape, and the cell-cavities are filled with a dark substance, described by Renault and Bertrand as protoplasm, and the cell-walls are fairly thick. In some of the larger specimens there are often found a few smaller sacs enclosed in the cavity of the partially disorganised mother-thallus. In the larger specimens the wall is usually invaginated in several places, giving the whole thallus a lobed or brain-like appearance. The supposed alga, which makes up ⁹⁄₁₀ths of the contents of a block of Kerosene shale, is named Reinschia Australis; it is regarded by the authors of the species as nearly related to the Hydrodictyaceae or Volvocineae.
In the Kerosene shale from certain localities in New South Wales Bertrand recognises a second form of thallus, which he refers to the genus Pila, characteristic of the European Bogheads.
The “thallus” characteristic of the Scotch Boghead has been named Pila scotica, and that of the Autun Boghead, Pila bibractensis.
In the latter form, which has been studied in more detail by MM. Renault and Bertrand, the thallus consists of about 6–700 cells, and is irregularly ellipsoidal in form, from ·189–·225mm. in length, and ·136–·160mm. broad. The surface-cells are radially disposed and pyramidal in shape, the internal cells are polygonal in outline and less regularly arranged (fig. 36, 2). The Pila thalli make up ¾ths of the mass in an average sample of the Autun Boghead. The Autun Boghead often contains siliceous nodules, and sections of these occasionally include cells of a Pila in which the protoplasmic contents and nuclei have been described by the French authors. The evidence for the existence of these supposed nuclei is, however, not entirely satisfactory; sections of silicified thalli which were shown to me by Prof. Bertrand did not satisfy me as to the minute histological details recognised by Bertrand and Renault.
The species of Pila are compared with the recent genus Celastrum, and regarded as most nearly allied to the Chroococcaceae or Pleurococcaceae among recent algae. Prof. Bornet[341] has suggested Gomphosphaeria as a genus which presents a resemblance to the Autun Pila.
In addition to the Bogheads of Autun, Torbanehill, and New South Wales, there are similar Palaeozoic deposits in Russia, America, and various other parts of the world. Full details of the structure of Boghead and the supposed algae referred to Reinschia, Pila, and other genera will be found in the writings of Bertrand and Renault[342].
The Kerosene shale of New South Wales affords the most striking and well-preserved examples of the cellular orange and yellow bodies referred to as the globular thalli of algae. It is almost impossible to conceive a purely inorganic material assuming such forms as those which occur in the Australian Boghead. On the other hand, it is hardly less easy to understand the possibility of such explanations as have been suggested of the organic origin of these characteristic bodies.
The ground-mass or matrix of the Boghead is referred to a brown ulmic precipitate thrown down on the floor of a Permian or Carboniferous lake, probably under the action of calcareous water. In this material there accumulated countless thalli of minute gelatinous algae, which probably at certain seasons completely covered the surface of the waters, as the fleurs d’eau in many of our fresh-water lakes. In addition to the thalli of Reinschia and Pila the Bogheads contain a few remains of various plant fragments, pollen-grains, and pieces of wood. Fish-scales and the coprolites of reptiles and fishes occur in some of the beds. On a piece of Kerosene shale in the Woodwardian Museum, Cambridge, there are two well-preserved graphitic impressions of the tongue-shaped fronds of Glossopteris Browniana, Brongn. There can be little doubt that the beds of Boghead were deposited under water as members of a regular sequence of sedimentary strata. The yellow bodies which form so great a part of the beds are practically all of the same type. Reinschia and Pila cannot always be distinguished, and it would seem that there are no adequate grounds for instituting two distinct genera and referring them to different families of recent algae.
Stated briefly, my conclusion is that the algae of the French authors may be definite organic bodies, but it is unwise to attempt to determine their affinities within such narrow limits as have been referred to in the above résumé. The structure of the bituminous deposits is worthy of careful study, and it is by no means impossible that further research might lead us to accept the view of the earlier investigators, that the brightly coloured organic-like bodies may be inorganic in origin.
D. RHODOPHYCEAE. (Florideae. Red Algae.)
The thallus of the members of this group assumes various forms, and consists of branched cell-filaments of a more or less complex structure. Cells of the thallus contain a red colouring matter in addition to the green chlorophyll. The reproduction is asexual and sexual; the formation of asexual reproductive cells (tetraspores) in groups of four in sporangia is a characteristic method of reproduction. Sexual reproduction is effected by means of distinct male and female cells.
With the exception of a few fresh-water genera all the red algae are marine. The Rhodophyceae, like the Cyanophyceae and Chlorophyceae, include a shell-boring form which has been found in the common razor-shell[343]. Several genera live as endophytes in the tissues of other algae. The recent species of this section of algae are characteristic of temperate and tropical seas. One subdivision of the red algae, the Corallinaceae, is extremely important from a geological point of view and must be dealt with in some detail.
Corallinaceae.
The thallus is usually encrusted with carbonate of lime; it is of a branched cylindrical form in the well-known Corallina officinalis, Linn. of the British coasts, of an encrusting and foliaceous type, in the genus Lithophyllum, and of a more coral-like form in the genus Lithothamnion. The reproductive organs occur in conceptacles, having the form of small depressed cavities in the thallus, or projecting as warty swellings above the surface of the plant. Asexual reproduction is by means of tetraspores formed in conceptacles resembling those containing the sexual cells. The Corallinaceae may be subdivided into the two families Melobesieae and Corallineae[344].
| Melobesieae. | Thallus encrusting, leaf- or coral-like; unsegmented. (Melobesia, Lithophyllum, Lithothamnion.) |
| Corallineae. | Cylindrical filamentous and segmented thallus. (Amphiroa and Corallina.) |
The genus Corallina is the best known British representative of the Corallinaceae. With other members of the group it was long regarded as a coralline animal, and it is only comparatively recently that the plant-nature of these forms has been generally admitted. Lithophyllum, Lithothamnion, Melobesia, and other genera of the Corallinaceae and some of the Siphoneae play a very important part in the building and cementing of coral-reefs. The pink or rose-coloured calcareous thallus of some of these calcareous algae or Nullipores imparts to coral-reefs a characteristic appearance. In some cases, indeed, the coral-reefs are very largely composed of algae. Saville Kent[345] describes the Corallines or Nullipores of the Australian Barrier-reef as furnishing a considerable quota towards the composition of the coral rock. Mr Stanley Gardiner, who accompanied the coral-boring expedition to the island of Funafuti, has kindly allowed me to quote the following extract from his notes, which affords an interesting example of the importance of calcareous algae as reef-building organisms. “It is quite a misnomer to speak of the outer edge of a reef like this (Rotuma Island) as being formed of coral. It would be far better to call it a Nullipore reef, as it is completely encrusted by these algae, while outside in the perfectly clear water, 10 to 15 fathoms in depth, the bottom has a most brilliant appearance from masses of red, white and pink Nullipores, with only a stray coral here and there.”
Agassiz[346] has given an account of the occurrence of immense masses of Nullipores (Udotea, Halimeda etc.) in the Florida reefs; his description is illustrated by good figures of these algae.
In the Mediterranean there are true Nullipore reefs, which are interesting geologically as well as botanically. Walther[347] has described one of these limestone-banks in the Gulf of Naples which occurs about 1 kilometre from the coast and 30 metres below the surface of the water. Every dredging, he says, brings up numberless masses of Lithothamnion fasciculatum (Lamarck), and L. crassum (Phil.). Between the branches of the algae, gasteropods and other animals become completely enclosed by the growing plants, while diatoms, foraminifera, and other forms of life are abundant. Water percolating through the mass gradually destroys the structure of the algal thalli, and in places reduces the whole bank to a compact structureless limestone.
The same author[348] has also called attention to the importance of Lithophyllum as a constructive element in the coral-reefs off the Sinai peninsula.
Lithothamnion a typical genus of the Corallinaceae may be briefly described.
Philippi[349] was the first writer to describe this and other genera as plants. He gave the following definition of Lithothamnion:
“Stirps calcarea rigida, e ramis cylindricis vel compressiusculis dichotoma ramosis constans.”
The thallus of Lithothamnion grows attached to the face of a rock or other foundation, and forms a hard, stony mass, assuming various coralline shapes. The exposed face may have the form of numerous short branches or of an irregular warty surface.
In section (fig. 37, A.) the lower part of the thallus is seen to be made up of rows of cells radiating out from a central point, and the upper portion consists of vertical and horizontal rows of cells. The whole body is divided up into a large number of small cells by anticlinal and periclinal walls, and possesses an evident cellular as distinct from a tubular structure. Conceptacles containing reproductive organs are either sunk in the thallus or project above the surface. The two types of structure in a single thallus are shown in fig. 37, A, also a conceptacle containing tetraspores.
In the closely allied Lithophyllum the thallus is encrusting, and in section it presents the same appearance as the lower part of a Lithothamnion thallus.
Species of Lithothamnion occur in the Mediterranean Sea, and are abundant in the arctic regions[353], while on the British coasts the genus is represented by four species[354]. Some large specimens of Lithothamnion and Lithophyllum are exhibited in one of the show-cases in the botanical department of the British Museum. For the best figures and descriptions of recent species reference should be made to the works of Hauck, Rosanoff, Rosenvinge, Kjellman and Solms-Laubach[355].
It is to be expected that such calcareous algae as Lithothamnion should be widely represented by fossil forms. In addition to the botanical importance of the data furnished by the fossil species as to the past history of the Corallinaceae, there is much of geological interest to be learnt from a study of the manner of occurrence of both the fossil and recent representatives. As agents of rock-building the coralline algae are especially important. The late Prof. Unger[356] in 1858 gave an account of the so-called Leithakalk of the Tertiary Vienna basin, and recognised the importance of fossil algae as rock-forming organisms. The Miocene Leithakalk, which is widely used in Vienna as a building stone[357], consists in part of limestone rocks consisting to a large extent of Lithothamnion.
Since the publication of Unger’s work several writers have described numerous fossil species of Lithothamnion from various geological horizons. A few examples will suffice to illustrate the range and structure of this and other genera of the Corallinaceae. In dealing with the fossil species it is often impossible to make use of those characters which are of primary importance in the recognition of recent species. The fossil thallus is usually too intimately associated with the surrounding rock to admit of any use being made of external form as a diagnostic feature. The size and form of the cells must be taken as the chief basis on which to determine specific differences. In the absence of conceptacles or reproductive organs it is not always easy to distinguish calcareous algae from fossil Hydrozoa or Bryozoa. In many instances, however, apart from the nature and size of the elements composing the thallus, the conceptacles afford a valuable aid to identification. An example of a fossil conceptacle containing tetraspores is shown in fig. 37, C; it is from a Tertiary species of Lithothamnion, described by Früh from Montévraz in Switzerland.
1. Lithothamnion mamillosum Gümb. Fig. 32, A (i) and (ii). (p. 155.) This species was first recorded by Gümbel[358] from the Upper Cretaceous (Danian) rocks of Petersbergs, near Maëstricht, on the Belgian frontier. It was originally described as a Bryozoan. The thallus has the form of an encrusting calcareous structure bearing on its upper surface thick nodular branches, as shown in fig. 32, A (ii); in section, A (i), the thallus consists of a regular series of rectangular cells.
The specific name mamillosum has also been given to a recent species by Hauck[359], but probably in ignorance of the existence of Gümbel’s Cretaceous species.
2. Lithothamnion suganum Roth. Fig. 37, B. The section of this form given in fig. 37, B shows three oval conceptacles filled with crystalline material. The two lower conceptacles originally communicated with the surface of the thallus, but as in recent species the deeper portions of the algal body became covered over by additions to the surface, forming merely dead foundations for new and overlying living tissues.
The cells of the thallus have a breadth of 7–9µ, and a length of 9–12µ.
The specimen was obtained from a Lithothamnion bank, probably of Upper Oligocene age, in Val Sugana[360], in the Austrian Tyrol.
Numerous other species of Jurassic, Cretaceous and Tertiary age might be quoted, but the above may suffice to illustrate the general characters and mode of occurrence of the genus. It is important that the student should become familiar with the Lithothamnion and Lithophyllum types of thallus, in view of their frequent occurrence in crystalline limestone rocks and in such comparatively recent deposits as those of upraised coral-reefs. The coral-rock of Barbados and other West-Indian islands affords a good illustration of the manner of occurrence of fossil coralline algae in association with corals and other organisms[361].
In the fossil species of Lithothamnion hitherto recorded there do not appear to be any important features in which they differ from recent forms; the geological history of the genus so far as it is known, favours the view that the generic characters are of considerable antiquity.
Mr A. Brown[362], of Aberdeen, has recently brought forward good evidence for including various calcareous fossils, described by several authors under different names and referred to various genera of fossil animals, in the genus Solenopora, which he places among the coralline algae.
Species of this genus have been described from England, Scotland, Esthonia, Russia, and other countries. The geological range of Solenopora appears to be from Ordovician to Jurassic rocks; in some cases it is an important constituent of beds of limestone.
Solenopora compacta (Billings). Fig. 38. This species was originally described by Billings as Stromatopora compacta, and afterwards defined by Nicholson and Etheridge. The thallus forms sub-spheroidal masses, from the size of a hemp-seed to that of an orange. The external surface is lobulate; the fractured surface has a porcellaneous and sometimes a fibrous appearance, and is usually white or light brown in colour. In vertical section (fig. 38, B) the cells are elongated and arranged in a radiating and parallel fashion; they often occur in concentric layers. The cells have a diameter of about ¹⁄₁₇ mm. and possess distinctly undulating walls, as seen in a tangential section (fig. 38, A). Brown describes certain larger cells in the thallus (fig. 38, A) as sporangia[363], but it is difficult to recognise any distinct sporangial cavities in the drawing. The example figured is from the Trenton limestone of Canada; a variety of the same species has been recorded from the Ordovician rocks of Girvan in Ayrshire. There appear to be good reasons for accepting Brown’s conclusion that Solenopora belongs to the Corallinaceae rather than to the Hydrozoa, among which it was originally included. After comparing Solenopora with recent genera of Florideae, Brown concludes that “the forms of the cells and cell-walls, the method of increase, and the arrangement of the tissue cells in the various species of Solenopora bear strong evidence of relationship between that genus and the calcareous algae[364].”
The importance of the calcareous Rhodophyceae has been frequently emphasised by recent researches, and our knowledge of the rock-building forms is already fairly extensive. We possess evidence of the existence of species of different genera in Ordovician seas, as well as in those of the Silurian, Triassic, Jurassic, and more recent periods. It is reasonable to prophesy that further researches into the structure of ancient limestones will considerably extend our knowledge of the geological and botanical history of the Corallinaceae.
Numerous fossils have been described as examples of other genera[365] of Rhodophyceae than those included in the Corallinaceae, but these possess little or no scientific value and need not be considered.
E. PHAEOPHYCEAE (Brown Algae).
Olive-brown algae, thallus often leathery in texture, composed of cell-filaments or parenchymatous tissue, in some cases exhibiting a considerable degree of internal differentiation. The sexual reproductive organs may be either in the form of passive egg-cells and motile antherozoids or of motile cells showing no external sexual difference.
With one or two exceptions all the genera are marine. They have a wide distribution at the present day, and are especially characteristic of far northern and extreme southern latitudes. The gigantic forms Lessonia, Macrocystis and others already alluded to, belong to this group; also the genus Sargassum, of which the numberless floating plants constitute the characteristic vegetation of the Sargasso Sea.
Palaeobotanical literature is full of descriptions of supposed fossil representatives of the brown algae, but only a few of the recorded species possess more than a very doubtful value; most of them are worthless as trustworthy botanical records. Many of the numerous impressions referred to as species of Fucoides and other genera present a superficial resemblance to the thallus of the common Bladder-wrack and other brown seaweeds. Such similarity of form, however, in the case of flat and branched algal-like fossils is of no scientific value. In many instances the impressions are probably those of an alga, but they are of no botanical interest. The flat and forked type of thallus of Fucus, Chondrus crispus (L.) and other members of the Phaeophyceae is met with also among the red and green algae, to say nothing of its occurrence in the group of thalloid Liverworts, or of the almost identical form of various members of the animal kingdom. The variety of form of the thallus in one species is well illustrated by the common Chondrus crispus (L.). This alga was described by Turner[366] in his classic work on the Fuci under the name of Fucus crispus as “a marine Proteus.” It affords an interesting example of the different appearance presented by the same species under different conditions, and at the same time it furnishes another proof of the futility of relying on imperfectly preserved external features as taxonomic characters of primary importance.
An example of a supposed Jurassic Fucus is shown in fig. 49, and briefly described in the Chapter dealing with fossil Bryophytes.
Several species of Flysch Algae have recently been referred by Rothpletz[367] to the Phaeophyceae under the provisional generic name Phycopsis, but they are of no special botanical interest.
The extremely interesting genus Nematophycus has lately been assigned by a Canadian author[368] to a position in the Phaeophyceae. Although the particular points on which he chiefly relies are not perhaps thoroughly established, there are certain considerations which lead us to include Nematophycus as a doubtful member of the present group of algae.
Nematophycus.
The stem attains a diameter of between 2 and 3 feet in the largest specimens; it is made up either of comparatively wide and loosely arranged tubes pursuing a slightly irregular vertical course accompanied by a plexus of much narrower tubes, or of tubes varying in diameter but not divisible into two distinct types. Rings of growth occur in some forms but not in others. Radially elongated or isodiametric spaces occur in the stem tissues in which the tubes are less abundant.
Reproductive organs unknown, with the possible exception of some very doubtful bodies described as spores.
In 1856 Sir William Dawson proposed the generic name Prototaxites for some large silicified trunks discovered in the Lower and Middle Devonian rocks of Canada. A few years later the same writer[369] published a detailed account of the new fossils and arrived at the conclusion that the Devonian stem showed definite points of affinity with the recent genus Taxus, and the generic name suggests that he regarded it as the type of Coniferous trees belonging to the sub-family Taxineae. The reasons for this determination were afterwards shown by Carruthers to be erroneous. Dawson thought he recognised pits and spiral thickenings in the walls of the tubular elements, as well as pointed ends in some of the latter. The spiral markings were in reality small hyphal tubes passing obliquely across the face of the wider tubes, and the apparent ends of the supposed tracheids were deceptive appearances due to the fact that the tubes had in some cases been cut through in an oblique direction. In 1870 Carruthers[370] expressed the opinion that Dawson’s Prototaxites was a “colossal fossil seaweed” and not a coniferous plant. The same author[371] in 1872 published a full and able account of the genus, and conclusively proved that Prototaxites could not be accepted as a Phanerogam; he brought forward almost convincing evidence in favour of including the genus among the algae. The name Prototaxites was now changed for that of Nematophycus. Carruthers compares the rings of growth in the fossil stems with those in the large Antarctic Lessonia stems, but he regards the histological characters as pointing to the Siphoneae as the most likely group of recent algae in which to include the Palaeozoic genus.
We may pass over various notes and additional contributions by Dawson, who did not admit the corrections to his original descriptions which Carruthers’ work supplied. In 1889 an important memoir appeared by Penhallow[372] of Montreal in which he confirmed Carruthers’ decision as to the algal nature of Prototaxites; he contributed some new facts to the previous account by Carruthers, and expressed himself in favour of regarding the fossil plant as a near ally of the recent Laminariae. The next addition to our botanical knowledge of this genus was made by Barber[373] who described a new specific type of Nematophycus—N. Storriei—found by Storrie in beds of Wenlock limestone age near Cardiff. Solms-Laubach[374], in a recent memoir on Devonian plants, recorded the occurrence of another species of this genus in Middle Devonian rocks near Gräfrath on the Lower Rhine. Lastly Penhallow[375], in describing a new species, lays stress on the resemblance of some of the tubular elements in the stem to the sieve-hyphae of the recent seaweeds Macrocystis and Laminaria. He concludes that the new facts he records make it clear that Nematophycus “is an alga, and of an alliance with the Laminarias.” The recent evidence brought forward by Penhallow is not entirely satisfactory; the drawings and descriptions of the supposed trumpet-shaped sieve-hyphae are not conclusive. On the whole it is probably the better course to speak of Nematophycus as a possible ally of the brown algae rather than as an extinct type of the Siphoneae, but until our knowledge is more complete it is practically impossible to decide the exact position of this Siluro-Devonian genus.
Solms-Laubach[376] has suggested that the generic name Nematophyton, used by Penhallow in preference to Carruthers’ term Nematophycus, is the more suitable as being a neutral designation and not one which assumes a definite botanical position. In view of the nature of the evidence in favour of the algal affinities of the fossil, the reasons for discarding Carruthers’ original name are hardly sufficient.
Before discussing more fully the distribution and botanical position of Nematophycus we may describe at length one of the best known species, and give a short account of some other forms.