Fossil plants, Vol. 1

1. Nematophycus Logani (Daws.). Fig. 39, A–E. The stem possesses well marked concentric rings of growth due to a periodic difference in size of the large tubular elements. The tissues consist of two distinct kinds of tubular elements, the larger tubes loosely arranged and pursuing a fairly regular longitudinal course, and having a diameter of 13–35µ; the smaller tubes, with a diameter of 5–6µ, ramify in different directions and form a loose plexus among the larger and more regularly disposed elements. Branching occurs in both kinds of tubes; septa have been recognised only in the smaller tubes. Irregular and discontinuous radial spaces traverse the stem tissues, having a superficial resemblance in their manner of occurrence to the medullary rays of the higher plants.

The best specimens of this species were obtained by Sir William Dawson from the Devonian Sandstones of Gaspé in New Brunswick. The largest stems had a diameter of 3 feet and reached a length of several feet[377]; in some examples Dawson found lateral appendages attached to the stem which he described as “spreading roots.” Externally the specimens were occasionally covered with a layer of friable coal, and internally the tissues were found to be more or less perfectly preserved by the infiltration of a siliceous solution. Most of the examples of Nematophycus from Britain and Germany are much smaller and less perfectly preserved than those from Canada. The Peter Redpath Museum, Montreal, contains several very large blocks of Nematophycus, in many of which one sees the concentric rings of growth clearly etched out by weathering agents in a cross section of a large stem.

In fig. 39, A, a sketch is given of a thin transverse section of a stem, drawn natural size. The lines of growth are clearly seen, and as in coniferous stems the breadth of the concentric zones varies considerably. The short lines traversing the tissues in a radial direction represent the medullary-ray-like spaces referred to in the specific diagnosis. A transverse section examined under a low-power objective presents the appearance of a number of thick-walled and comparatively wide tubes loosely arranged; they may be in contact or separated from one another. If the microscope be carefully focussed through the thickness of the section the transversely-cut tubes appear to move laterally, producing a curiously dazzling effect if the objective is raised or lowered rapidly. This lateral movement is due to the undulating vertical course of the tubes. Under a higher power the lighter-coloured matrix in which the tubes are embedded shows a number of very much smaller and thinner-walled hyphal elements; some of these are cut across transversely, others more or less obliquely and others again longitudinally. These smaller tubes constitute an irregular plexus surrounding and ramifying between the larger elements. The diameter of the larger tubes decreases for a certain distance in a radial direction as seen in a transverse section, and this change in size gives rise to the appearance of concentric lines indicating periodic changes in growth.

The radial spaces are characterised by the partial absence of the larger tubes, and as seen in longitudinal sections these spaces constitute regions in which the smaller tubes branch very freely. Fig. 39, B, represents a small piece of a transverse section seen under a fairly high power. In fig. 39, C, the tubes are seen in longitudinal section. The larger elements are unseptate and not very regular in their vertical course through the stem; the smaller elements are seen as fine tubes lying between and across the larger tubes. In the sections I have examined no undoubted transverse septa were detected in any of the tubular elements.

The question as to the possible connection between the larger and smaller elements is one which is not as yet satisfactorily disposed of. Penhallow[378] regards the finer hyphal elements as branches of the larger tubes, but Barber[379], who has carefully examined good material of Nematophycus Logani, was unable to detect any organic connection between the two. My own observations are in accord with those of Barber. Further details and numerous figures of this species of Nematophycus will be found in the memoirs of Carruthers, Penhallow and Barber.

Some specimens of silicified Nematophycus stems afford particularly instructive examples of the state of preservation or method of mineralisation as a source of error in histological work. The sketches reproduced in fig. 39, D and E, were made from a section of a large specimen of Nematophycus in the British Museum. In fig. D we have one of the radial spaces containing some indistinct small elements, the tissue surrounding the space appears to consist of polygonal cells suggesting ordinary parenchymatous tissue. In fig. E a few of these ‘cells’ are seen more clearly, they have black and ragged walls, and often contain very small and faint circles of which the precise nature is uncertain. The true interpretation of this form of structure was first supplied by Penhallow[380]. The black network simulating parenchymatous tissue consists of the substance of Nematophycus tubes which has been completely redistributed during fossilisation and collected along fairly regular lines, as seen in figs. D and E. The original structure has been almost completely destroyed, and the material composing the walls of the large tubes has finally been rearranged as a network, interrupted here and there by the characteristic radial spaces which remain as evidence of the original Nematophycus characters. It is possible in some cases to trace every gradation from sections exhibiting the normal structure through those having the appearance shown in figs. D and E to others in which the structure is completely lost. Penhallow describes this method of fossilisation in N. crassus (Daws.); an examination of several specimens in the National Collection leads me to entirely confirm his general conclusions, and also to the opinion that N. Logani shows exactly the same manner of mineralisation as N. crassus. The chief point of interest as regards this method of preservation lies in the fact that a fossil described by Dawson[381] as Celluloxylon primaevum, and referred to as a probable conifer, is undoubtedly a badly preserved Nematophycus. Penhallow examined Dawson’s specimens and obtained convincing evidence of their identity with certain forms of highly altered Nematophycus stems.

2. Nematophycus Storriei Barber. Fig. 40. The specimens on which Barber[382] founded this species were obtained by Mr Storrie from the Tymawr quarry near Cardiff, in beds of Wenlock age. The fragmentary nature of the material is largely compensated for by the excellence of the preservation. We may briefly define the species as follows:

The stem consists of separate interlacing undivided and usually unbranched tubes of varying diameter. Spaces more or less isodiametric in dimensions are scattered through the tissue. The spaces constitute regions in which the tubular elements branch freely.

The main distinguishing features of this British species are (i) the absence of two distinct and well-defined forms of tubular elements. The main part of the stem consists of thick walled tubes similar to those of N. Logani, but the spaces between them are occupied by thinner-walled and smaller tubes varying considerably in diameter; (ii) the form of the spaces which are not radially elongated as in N. Logani.

Fig. 40 shows the undulating course of the tubes as seen in a longitudinal section; the black colour of some of the elements is due to the fact that the surface of the wall is seen, while in the lighter-coloured portions of the tubes the wall has been cut through. The lighter patch about the middle of the figure shows the form of one of the spaces in which the tubes are freely branched.

In addition to the two species already described six others have been recorded, but with these we need not concern ourselves in detail. One of these species, N. Hicksi, was found by Dr Hicks[383] in the Denbighshire grits quarry of Pen-y-Glog near Corwen in North Wales. The position of these beds has recently been determined by Mr Lake[384] as corresponding to that of the Wenlock limestone. This species and N. Storriei are both Silurian examples of the genus. It is possible, as Barber has suggested, that the specimens described under these two names should be referred to one species. The specimens found by Hicks were small and imperfectly preserved fragments; Etheridge has given a full description of their structure, and Barber has subsequently examined the material. The preservation is not such as will admit of any very precise specific diagnosis; the fragments are correctly referred to Nematophycus, but their specific characters cannot be clearly determined.

Solms-Laubach[385] has described some fragments of another species of Nematophycus from the Devonian rocks of the Lower Rhine. His specimens are chiefly interesting as extending the geographical range of the genus, and as affording examples of a curious method of preservation. The specimens obtained were small fragments, flattened and very dark brown in colour. The tubular elements consisted of an external membrane of black coal, enclosing a central core of dark red iron-oxide. On burning the fragment on a piece of platinum foil the coal composing the wall of the tubes was removed and the deep-red casts of the tube-cavities remained[386]. The investigation of the structural characters of this imperfect material was conducted by reflected light. Under certain conditions, when it is impossible to obtain thin sections for examination by transmitted light, it is possible to accomplish much, as shown by Solms-Laubach’s work, by means of observation with direct light.

The last species to be noticed is Nematophycus Ortoni recently described by Penhallow. There are no concentric rings of growth, no radial spaces and no smaller hyphae in the tissues of this type of stem. In longitudinal section, the tubes show occasional local expansions of the lumen which Penhallow compares with the ‘trumpet-hyphae’ of some recent brown algae. No actual sieve-plates or transverse walls have been detected, but the general appearance of the tubes is considered to afford distinct evidence of the original existence of such walls. The figures accompanying the description do not carry conviction as to the correctness of the reference of the tubes to imperfectly preserved sieve-hyphae.

The following list, taken, with a few alterations, from Penhallow’s memoir[387], shows the geographical and geological range of the species of Nematophycus hitherto recorded.

In summing up our information as to the structure of Nematophycus we find there are certain points not definitely settled, and which are of considerable importance. The few recorded instances of spore-like bodies by Penhallow and Barber are not satisfactory; we are still ignorant of the nature of the reproductive organs. Such instances of lateral appendages as have been referred to do not throw much light on the habit of the plant. So far as we know at present the stem of Nematophycus was not differentiated internally into a cortical and central region. It may be that the specimens have been only partially preserved, and the coaly layer which occasionally surrounds a stem may represent a carbonised cortex which has never been petrified. The large and loosely arranged tubes constitute the chief characteristic feature of the genus; in some cases (N. Logani) there is an accompanying plexus of smaller hyphae, in others (N. Storriei) there is no definite division of the tissue into two sets of tubes of uniform size, and in N. Ortoni the tubular elements are all of the large type.

Penhallow has recognised the branching of large tubes in N. Logani and N. crassus giving rise to the small hyphal elements. In most specimens, however, no such mode of origin of the smaller tubes can be detected. The spaces which interrupt the homogeneity of the tissues in some forms have been described as branching depots, on account of the frequent occurrence in these areas of much branched hyphae. The function of these spaces (fig. 39, D, and fig. 40) may be connected with aeration of the stem-tissues.

As Carruthers first pointed out the unseptate nature of the elements and the occurrence of large and small tubes forming a comparatively lax tissue suggested affinities with such recent genera as Penicillus, Halimeda, Udotea and other members of the Siphoneae. In those fossil stems which possess tubes of two distinct sizes, we cannot as a rule trace any organic connection between the two sets of tubular elements. Transverse septa have been detected in the tubes of some specimens of N. Logani. These considerations and the large size and habit of growth of the stem leave one sceptical as to the wisdom of assigning the fossil genus to the Siphoneae. On the other hand, apart from the doubtful sieve-hyphae of Penhallow, the manner of growth of the plant, the concentric rings, marked by a decrease in the diameter of the tubes, the lax arrangement and irregular course of the elements, afford points of agreement with some recent Phaeophyceae. The stem of a Laminaria (fig. 29) or of a Lessonia are the most obvious structures with which to compare Nematophycus. The medullary region of a Laminaria or Fucus and of other genera presents a certain resemblance to the tissues of the fossil stems. On the whole we may be content to leave Nematophycus for the present as probably an extinct type of alga, more closely allied to the large members of the Phaeophyceae than to any other recent seaweeds.

Pachytheca.

There is another fossil occasionally associated with Nematophycus and referred by many writers to the Algae, which calls for a brief notice. Pachytheca is too doubtful a genus to justify a detailed treatment in the present work. Although, as I have elsewhere suggested[389], we are hardly in a position to speak with any degree of certainty as to its affinity, it is not improbable that it may eventually be shown to be an alga.

Without attempting a full diagnosis of the genus, we may briefly refer to its most striking characters.

Pachytheca usually occurs in the form of small spherical bodies, about ·5 cm. in diameter, in Old Red Sandstone or Silurian rocks. In section a single sphere is found to consist of two well marked regions; in the centre, of a number of ramifying and irregularly placed narrow tubes, and in the peripheral or cortical region, of numerous regular and radially disposed simple or forked septate tubes. The tubular elements of the two regions are in organic connection.

The name was proposed by Sir Joseph Hooker for some specimens found by Dr Strickland[390] in the Ludlow bone-bed (Silurian) of Woolhope and May-Hill. Examples were subsequently recorded from the Wenlock limestone of Malvern and from Silurian and Old Red Sandstone rocks of other districts. Hicks[391] found Pachytheca in the Pen-y-Glog grits of Corwen in association with Nematophycus, and the two fossils have been found together elsewhere. This association led to the suggestion that Pachytheca might be the sporangium of Nematophycus, and Dawson[392], in conformity with his belief in the coniferous character of the latter plant, referred to Pachytheca as a true seed.

The best sections of this fossil have been prepared with remarkable skill by Mr Storrie of Cardiff; they were carefully examined and described by Barber in two memoirs[393] published in the Annals of Botany, the account being illustrated by several well executed drawings and microphotographs.

Among other difficulties to contend against in the interpretation of Pachytheca there is that of mineralisation. The preservation is such as to render the discrimination of original structure as distinct from structural features of secondary origin, consequent on a particular manner of crystallisation of the siliceous material, a matter of considerable difficulty.

Suggestions as to the nature of Pachytheca have been particularly numerous; it has been referred to most classes of plants and relegated by some writers to the animal kingdom. The most recent addition to our knowledge of this problematic fossil was the discovery of a specimen by Mr Storrie in which the Pachytheca sphere rested in a small cup, like an acorn fruit in its cupule. This specimen was figured and described by Mr George Murray[394] in 1895; he expresses the opinion that the discovery makes the taxonomic position of the genus still more obscure. Solms-Laubach briefly refers to Pachytheca in connection with Nematophycus, and regards its precise nature almost as much an unsolved riddle now as it was when first discovered. For a fuller account of this fossil reference must be made to the contributions of Hooker[395], Barber[396] and others. The literature is quoted by Barber and more recently by Solms-Laubach[397]. There are several specimens and microscopic sections of Pachytheca in the geological and botanical departments of the British Museum. The genus has been recorded from Shropshire, North Wales, Malvern, Herefordshire, Perthshire and other British localities, as well as from Canada; it occurs in both Silurian and Old Red Sandstone rocks.

Algites.

A generic name for those fossils which in all probability belong to the class Algae, but which, by reason of the absence of reproductive organs, internal structure, or characters of a trustworthy nature in the determination of affinity, cannot be referred with any degree of certainty to a particular recent genus or family.

This term was suggested in 1894[398] as a provisional and comprehensive designation under which might be included such impressions or casts as might reasonably be referred to Algae. The practice of applying to alga-like fossils names suggestive of a definite alliance with recent genera is as a rule unsound. It would simplify nomenclature, and avoid the multiplication of generic names, if the term Algites were applied to such algal fossils from rocks of various ages as afford no trustworthy data by which their family or generic affinity can be established.

V. MYXOMYCETES (MYCETOZOA).

This class of organisms affords an interesting example of the impossibility of maintaining a hard and fast line between the animal and plant kingdom. Zoologists and Botanists usually include the Myxomycetes[399] in the text-books of their respective subjects, and the name Animal-fungi which has been applied to these organisms expresses their dual relationship. They constitute one of three groups which we may include in that intermediate zone or ‘buffer-state’ between the two kingdoms. From a palaeobotanical point of view the Myxomycetes are of little interest, but a very brief reference may be made to them rather for the sake of avoiding unnecessary incompleteness in our classification than from their importance as possible fossils.

They are organisms without chlorophyll, consisting of a naked mass of protoplasm, known as the plasmodium, which may attain a size of several inches. Such plasmodia creep over the surface of decaying organic substrata, and in forming their asexual reproductive cells they are converted into somewhat complex fruits containing spores. The spores produce motile swarm-cells, which eventually coalesce together to form a new plasmodium.

A few examples of fossil Myxomycetes have been recorded from the Palaeozoic and more recent formations, but none of them are entirely beyond suspicion. We may mention three examples of fossils referred to this group, but only one of these is entitled to serious consideration.

Myxomycetes Mangini Ren.[400] It is not uncommon to find distinct traces of original or secondary cell-contents in well preserved petrified plant-tissues. There is often a difficulty, however, in distinguishing between the true cell-contents and the cells of some parasitic or saprophytic intruder. In some petrified corky tissue in a silicified nodule from the Permo-Carboniferous beds of Autun, Renault has recently discovered what he believes to be traces of a Myxomycetous plasmodium. The cork-cells would be without protoplasmic contents of their own, and their cavities contain a number of fine strands stretching from the cell-walls in different directions and uniting in places as irregular or more or less spherical masses. The drawings given by Renault of these irregular reticulated structures with scattered patches of what may possibly be petrified plasmodial protoplasm bear a striking resemblance to the plasmodium of a Myxomycete. A figure of the capillitium of a species of Leocarpus figured by Schröter[401] in his account of the Myxomycetes in Engler and Prantl’s work is very similar to that of Renault’s ‘plasmodium.’

It is by no means inconceivable that the Myxomycetes Mangini may be correctly referred to this group, but the wisdom of assigning a name to such structures may well be questioned.

The other two examples call for little notice. Messrs Cash and Hick[402] in a paper on fossil fungi from the Coal-Measures refer to some small spherical bodies as possibly the spores of a Myxomycete. They might be referred equally well to numerous other organisms.

Göppert and Menge[403] in their monograph on plants in the Baltic Tertiary Amber, express the opinion that an ill-defined tangle of threads which they figure may be a Myxomycete.

It would serve no useful purpose to quote other instances of possible representatives of fossil Mycetozoa; but the consideration of the above examples may serve to emphasize the desirability of refraining from converting a possibility into an apparently recognised fact by the application of definite generic and specific names.

VI. FUNGI.

The most striking difference between the fungi and algae is the absence of chlorophyll in the former, and the consequent inability of fungi to manufacture their organic compounds from inorganic material. Fungi live therefore either as parasites or saprophytes, and as the same species may pass part of its life in a living host to occur at another stage of its development as a saprophyte, it is impossible to distinguish definitely between parasitic and saprophytic forms. The vegetative body of a fungus, that is the portion which is concerned with providing nourishment and preparing the plastic food-substance for the reproductive organs, is known as the mycelium. It consists either of a single and branched tubular cell known as a hypha, or of several hyphae or thread-like elements (filamentous fungi). The hyphal filaments may be closely packed together and form a felted mass of compact tissue, which in cross section closely simulates the parenchyma of the higher plants. This pseudoparenchymatous form of thallus is particularly well illustrated by the so-called sclerotia; these are sharply defined and often tuberous masses of hyphal tissue covered by a firm rind and containing supplies of food in the inner hyphae. They are able to remain in a quiescent state for some time, and to resist unfavourable conditions until germination and the formation of a new individual take place. The reproductive structures assume various forms; in some of the simpler fungi (Phycomycetes) sexual organs occur, as in the parallel group of Siphoneae among the algae, but in the higher fungi the reproduction is usually entirely asexual. An interesting case has recently been recorded among the more highly differentiated fungi in which distinct sexuality has been established[404]. In addition to the reproductive organs, such as oogonia and antheridia, the asexual cells or spores are borne either in special sporangia, or they occur as exposed conidia on supporting hyphae or conidiophores. Thick-walled and resistant resting-spores of various forms are also met with.

Without going into further details we may very briefly refer to the larger subdivisions of this group of Thallophytes.

Before describing a few examples of fossil fungi, it is important to consider the general question of their manner of occurrence and determination. Considering the small size and delicate nature of most fungi, it is not surprising that we have but few satisfactory records of well-defined fossil forms. The large leathery sporophores of Polyporus and other genera of the Basidiomycetes, which are familiar objects as yellow or brown brackets projecting from the trunks of diseased forest trees, have been found in a fairly perfect condition in the Cambridgeshire peat-beds, and examples have been described also by continental writers[405]. As a general rule, however, we have to depend on the chance mineralisation or petrifaction of the hyphae of a fungus-mycelium which has invaded the living or dead tissues of some higher plant. In the literature on fossil plants there are numerous recorded species of fungi founded on dark coloured spots and blotches on the impression of a leaf. Most of such records are worthless; the external features being usually too imperfect to allow of accurate identification. The occurrence of recent fungi as discolourations on leaves is exceedingly common, and the characteristic perithecia or compact and more or less spherical cases enclosing a group of sporangia in certain Ascomycetous species, might be readily preserved in a fossil condition.

Ascomycetes.

Some examples of possible Ascomycetous fungi have been recently recorded by Potonié from leaves and other portions of plants of Permian age. There is a distinct superficial resemblance between the specimens he figures and the fructifications of recent Ascomycetes, but in the absence of internal structure, it would be rash to do more than suggest the probable nature of the markings he describes. For one of the fungus-like impressions Potonié proposes the generic name Rosellinites; he compares certain irregularly shaped projections on a piece of Permian wood with the perithecia of Rosellinia, a member of the Sphaeriaceae, and describes them as Rosellinites Beyschlagii Pot.[406] Various other records of similar Ascomycetes-like fossils may be found in palaeobotanical literature[407], but it is unnecessary to examine these in detail. Unless we are able to determine the nature of the supposed fungus by microscopical methods our identifications cannot in most cases be of any great value.

An example of the perithecia of a fungus (Rosellinia congregata [Beck])[408] has been recorded from the Oligocene of Saxony, which would appear to rest on a more satisfactory basis than is often the case. In this particular instance the small projections on a piece of fossil coniferous stem present a form which naturally suggests a fungus perithecium. In cases where the black spots on a fossil stem or leaf possess a definite form and structure, it is perfectly legitimate to refer them to a group of fungi; but in very many instances the forms referred to such genera as Sphaerites and others are of little or no value. Many forms of scale-insects and galls on leaves present an obvious superficial resemblance to epiphyllous fungi, and might readily be mistaken for the fructifications of certain Ascomycetous species. As examples of scale-insects simulating fungi, reference may be made to such genera of the Coccineae as Aspidiotus, Diaspis, Lecanium, Coccus, and others. The female insects lying on the surface of a leaf, if preserved as a fossil impression, might easily be mistaken for perithecia[409].

Another pitfall in fossil mycology may be illustrated by a description of a supposed fungus, Sclerotites Salisburiae[410], Mass. on a Tertiary Ginkgo leaf. The figure given by Massalongo represents a Ginkgo leaf with well marked veins, the lamina between the veins being traversed by short discontinuous and longitudinally-running lines; the latter are referred to as the fungus. In a recent Ginkgo leaf one may easily detect with the naked eye a number of short lines between and parallel to the veins, which if examined in section are found to be secretory canals. There can be little doubt that Sclerotites Salisburiae owes its existence to the preservation of these canals.

The list of fossil fungi given by Meschinelli in Saccardo’s Sylloge Fungorum[411] includes certain species which are of no botanical value, and should have no place in any list which claims to be authentic.

Among the numerous examples of fossil ‘fungi’ which have no claim to be classed with plants, there are some which are in all probability the galleries of wood-eating animals. The radiating grooves frequently found on the inner face of the bark of a pine tree made by species of the beetle Bostrychus might be mistaken for the impressions of the firm strands of mycelial tissue of some Basidiomycetous fungus.

In some notes on fossil fungi by J. F. James[412] contributed to the American Journal of Mycology in 1893, it is pointed out that a supposed fungus described by Lesquereux from the Lower Coal-Measures as Rhizomorpha Sigillariae[413], bears a strong likeness to some insect-burrows, such as those of Bostrychus.

“A new fungus from the Coal-Measures” described by Herzer in 1893[414] may probably be referred to animal agency. In any case there is no evidence as to the fungoid nature of the object represented in the figure accompanying Herzer’s description.

Basidiomycetes.

More trustworthy evidence of fossil fungi is afforded by the marks of disease in petrified tissue and by the presence of true mycelia. In examining closely the calcareous and siliceous plant-tissues from the Coal-Measures and other geological horizons, one occasionally sees fine thread-like hyphae ramifying through the cells or tracheal cavities; in many cases the hyphae bear no reproductive organs and cannot as a rule be referred to a particular type of fungus. If the hyphal filaments are unseptate, they most likely belong to some Phycomycetous species; or if they are obviously septate the Mesomycetes or the Mycomycetes are the more probable groups. Occasionally there may be found indications of the characteristic clamp-connections in the septate filaments; a small semicircular branch, which is given off from a mycelium immediately above a transverse wall, bends round to fuse with the filament just below the septum, thus serving as a small loop-line connecting the cell-cavity above and below a cross wall. Such clamp-connections are usually confined to the hyphae of Basidiomycetes and thus serve as a useful aid in identification. A good example of a clamp-connection in a fossil mycelium is figured by Conwentz[415] in his monograph on the Baltic amber-trees of Oligocene age. The stout and thick type of hypha found in some fossil woods agrees closely with that of Polyporus, Agaricus melleus and other well-known recent Basidiomycetes.

In a section of a piece of lignified coniferous wood recently brought by Col. Feilden from Kolguev island[416], the brown and stout hyphae of a fungus are clearly seen as distinct dark lines traversing the tracheal tissue. The occurrence of septa and the large diameter of the mycelial branches at once suggest a comparison with such recent forms as Agaricus melleus, Polyporus and other Basidiomycetes. The age of the Kolguev wood is not known with any certainty.

The vesicular swellings such as those represented in fig. 41, A, B, D and E, may easily be misinterpreted. Such spherical expansions in a mycelium, either terminal or intercalary, may be sporangia, oogonia or large resting-spores, or non-fungal cell-contents, and it is usually impossible in the absence of the contents to determine their precise nature. Hartig[417] and others have drawn attention to the occurrence of such bladder-like swellings in the mycelia of recent fungi, which have nothing to do with reproductive purposes; under certain conditions the hyphae of a fungus growing in the cavity of a cell or trachea may form such vesicles, and these, as in fig. 42, D, m may completely fill up the cavity of a large tracheid.

Some good examples of bladder-like swellings, such as occur in the mycelium of Agaricus melleus and other recent fungi, have been figured by Conwentz[418] in fossil wood of Tertiary age from Karlsdorf. The swellings in this fossil fungus might easily be mistaken for oogonia or sporangia; especially as they are few in number and spherical in form.

A similar appearance is presented by a mass of tyloses in the cavity of an old vessel or tracheid; and vesicular cell-contents, as in the cells of fig. 41, A, 2–5, may closely simulate a number of thin-walled fungal spores or sporangia.

A good example of such a vesicular tissue, in addition to that already quoted, is afforded by a specimen of an Eocene fern, Osmundites Dowkeri Carr.[419] described by Carruthers in 1870. The ground-tissue cells contain traces of distinct fungal hyphae (fig. 41, B), and in many of the parenchymatous elements the cavity is completely filled with spherical vesicles; in other cases one finds hyphae in the centre of the cell while vesicles line the wall, as shewn in fig. 41, B. Carruthers refers to these bladders as starch grains, and this may be their true nature; their appearance and abundant occurrence in the parenchyma certainly suggest vesicular cell-contents rather than fungal cells. I could detect no proof of any connection between the hyphae and bladders, and the absence of the latter in the cavities of the tracheids, fig. 41, C, favoured the view of their being either starch-grains or other vacuolated contents similar to that in the cells of the Portland Cycad (fig. 41, A) referred to on p. 88.

The vacuolated cell-contents partially filling the cells in fig. 41, D, present a striking resemblance to the contents of the cells 2–5 in fig. 41, A. In fig. D the frothy and contracted substance might be easily mistaken for a parasitic or saprophytic fungus, but this resemblance is entirely misleading. It is by no means uncommon to find the cells of recent plants occupied by such vacuolated contents, especially in diseased tissues in which a pathological effect produces an appearance which has more than once misled the most practised observers.

In the important work recently published by Renault on the Permo-Carboniferous flora of Autun, there is a small spore-like body described as a teleutospore, and classed with the Puccineae[420]. We have as yet no satisfactory evidence of the existence of this section of Fungi in Palaeozoic times, and Renault’s description of Teleutospora Milloti from Autun might be seriously misleading if accepted without reference to his figure. The fragment he describes cannot be accepted as sufficient evidence for the existence of a Palaeozoic Puccinia.

The same author refers another Palaeozoic fungus to the Mucorineae under the name of Mucor Combrensis[421]; this identification is based on a mycelium having a resemblance to the branched thallus of Mucor, but in the absence of reproductive organs such resemblance is hardly adequate as a means of recognition.

The occurrence of hyphal cells in calcareous shells and corals has already been alluded to.[422] In addition to the examples referred to above, there is one which has been described by Etheridge[423] from a Permo-Carboniferous coral. This observer records the occurrence of tubular cavities in the calices of Stenopora crinita Lonsd., and attributes their origin to a fungus which he names Palaeoperone endophytica; he mentions one case in which a tube contains fine spherical spore-like bodies which he compares with the spores of a Saprolegnia. As pointed out above (p. 128), it is almost impossible to decide how far these tubes in shells and corals should be attributed to fungi, and how far to algae.