CHAPTER V.

DIFFICULTIES AND SOURCES OF ERROR IN THE DETERMINATION OF FOSSIL PLANTS.

“Robinson Crusoe did not feel bound to conclude, from the single human footprint which he saw in the sand, that the maker of the impression had only one leg.”
Huxley’s Hume, p. 105, 1879.

The student of palaeobotany has perhaps to face more than his due share of difficulties and fruitful sources of error; but on the other hand there is the compensating advantage that trustworthy conclusions arrived at possess a special value. While always on the alert for rational explanations of obscure phenomena by means of the analogy supplied by existing causes, and ready to draw from a wide knowledge of recent botany, in the interpretation of problems furnished by fossil plants, the palaeobotanist must be constantly alive to the necessity for cautious statement. That there is the greatest need of moderation and safe reasoning in dealing with the botanical problems of past ages, will be apparent to anyone possessing but a superficial acquaintance with fossil plant literature. The necessity for a botanical and geological training has already been referred to in a previous chapter.

It would serve no useful purpose, and would occupy no inconsiderable space, to refer at length to the numerous mistakes which have been committed by experienced writers on the subject of fossil plants. Laymen might find in such a list of blunders a mere comedy of errors, but the palaeobotanist must see in them serious warnings against dogmatic conclusions or expressions of opinion on imperfect data and insufficient evidence. The description of a fragment of a handle of a Wedgwood teapot as a curious form of Calamite[137] and similar instances of unusual determinations need not detain us as examples of instructive errors. The late Prof. Williamson has on more than one occasion expressed himself in no undecided manner as to the futility of attempting to determine specific forms among fossil plants, without the aid of internal structure[138]; and even in the case of well-preserved petrifactions he always refused to commit himself to definite specific diagnoses. In his remarks in this connection, Williamson no doubt allowed himself to express a much needed warning in too sweeping language. It is one of the most serious drawbacks in palaeobotanical researches that in the majority of cases the specimens of plants are both fragmentary and without any trace of internal structure. Specimens in which the anatomical characters have been preserved necessarily possess far greater value from the botanist’s point of view than those in which no such petrifaction has occurred. On the other hand, however, it is perfectly possible with due care to obtain trustworthy and valuable results from the examination of structureless casts and impressions. In dealing with the less promising forms of plant fossils, there is in the first place the danger of trusting to superficial resemblance. Hundreds of fossil plants have been described under the names of existing genera on the strength of a supposed agreement in external form; but such determinations are very frequently not only valueless but dangerously misleading. Unless the evidence is of the best, it is a serious mistake to make use of recent generic designations. If we consider the difficulties which would attend an attempt to determine the leaves, fragments of stems and other detached portions of various recent genera, we can better appreciate the greater probability of error in the case of imperfectly preserved fossil fragments.

EXTERNAL RESEMBLANCE.

The portions of stems represented in figures 20 and 21, exhibit a fairly close resemblance to one another; in the absence of microscopical sections or of the reproductive organs it would be practically impossible to discriminate with any certainty between fossil specimens of the plants shown in the drawings. Examples such as these, and many others which might be cited, serve to illustrate the possibility of confusion not merely between different genera of the same family, but even between members of different classes or groups. The long slender branches of the Polygonum represented in (fig. 21) would naturally be referred to Equisetum in the absence of the flowers (fig. 20 B), or without a careful examination of the insignificant scaly leaves borne at the nodes. The resemblance between Casuarina and Ephedra and the British species of Equisetum, or such a tropical form as E. debile, speaks for itself.

Fig. 20.

Restio tetraphylla Labill. (Monocotyledon).
Equisetum variegatum Schleich. (Vascular Cryptogam).
Equisetum debile Roxb. (Vascular Cryptogam)
Casuarina stricta Dryand. (Dicotyledon).
Ephedra distachya Linn. (Gymnosperm). (A–E ½ nat. size).

Fig. 21. Polygonum Equisetiforme Sibth. and Sm. A. Showing habit of plant. ½ nat. size. The two flowers towards the apex of one branch, drawn to a larger scale in B. C. Node with small leaf and ochrea characteristic of Polygonaceæ. From a plant in the Cambridge Botanic Garden.

Fig. 22. Kaulfussia æsculifolia Blume. From a specimen from Java in the British Museum herbarium. ⅓ nat. size.

Endless examples might be quoted illustrating the absolute futility, in many cases, of relying on external features even for the purpose of class distinction. An acquaintance with the general habit and appearance of only the better known members of a family, frequently leads to serious mistakes. The specimen shown in fig. 22 is a leaf of a tropical fern Kaulfussia, a genus now living in South-eastern Asia, and a member of one of the most important and interesting families of the Filicinæ, the Marrattiaceæ; its form is widely different from that which one is accustomed to associate with fern fronds. It is unlikely that the impression of a sterile leaf of Kaulfussia would be recognised as a portion of a fern plant.

Similarly in another exceedingly important group of plants, the Cycadaceæ[139], the examples usually met with in botanical gardens are quite insufficient as standards of comparison when we are dealing with fossil forms. Familiarity with a few commoner types leads us to regard them as typical for the whole family. In Mesozoic times cycadean plants were far more numerous and widely distributed than at the present time, and to adequately study the numerous fossil examples we need as thorough an acquaintance as possible with the comparatively small number of surviving genera and species. The less common and more isolated species of an existing family may often be of far greater importance to the palaeobotanist than the common and more typical forms. This importance of rare and little known types will be more fully illustrated in the chapters dealing with the Cycadaceæ and other plant groups. Among Dicotyledons, the Natural Order Proteaceæ, at present characteristic of South Africa and Australia, and also represented in South America and the Pacific Islands, is of considerable interest to the student of fossil Angiosperms. In a valuable address delivered before the Linnean Society[140] in 1870 Bentham drew attention to the marked ‘protean’ character of the members of this family. He laid special stress on this particular division of the Dicotyledons in view of certain far-reaching conclusions, which had been based on the occurrence in different parts of Europe of fossil leaves supposed to be those of Proteaceous genera[141]. Speaking of detached leaves, Bentham says:—“I do not know of a single one which, in outline or venation, is exclusively characteristic of the order, or of any one of its genera.” Species of Grevillea, Hakea and a few other genera are more or less familiar in plant houses, but the leaf-forms illustrated by the commoner members of the family convey no idea of the enormous variation which is met with not only in the family as a whole, but in the different species of the same genus. The striking diversity of leaf within the limits of a single genus will be dealt with more fully in volume II. under the head of Fossil Dicotyledons.

VENATION CHARACTERS.

There is a common source of danger in attempting to carry too far the venation characters as tests of affinity. The parallel venation of Monocotyledons is by no means a safe guide to follow in all cases as a distinguishing feature of this class of plants. In addition to such leaves as those of the Gymnosperm Cordaites and detached pinnæ of Cycads, there are certain species of Dicotyledons which correspond in the character of their venation to Monocotyledonous leaves. Eryngium montanum Coult., E. Lassauxi Dcne., and other species of this genus of Umbelliferæ agree closely with such a plant as Pandanus or other Monocotyledons; similarly the long linear leaves of Richea dracophylla, R. Br., one of the Ericaceæ, are identical in form with many monocotyledonous leaves. Instances might also be quoted of monocotyledonous leaves, such as species of Smilax and others which Lindley included in his family of Dictyogens which correspond closely with some types of Dicotyledons[142]. Venation characters must be used with care even in determining classes or groups, and with still greater reserve if relied on as family or generic tests.

It is too frequently the case that while we are conversant with the most detailed histological structure of a fossil plant stem, its external form is a matter of conjecture. The conditions which have favoured the petrifaction of plant tissues have as a rule not been favourable for the preservation of good casts or impressions of the external features; and, on the other hand, in the best impressions of fern fronds or other plants, in which the finest veins are clearly marked, there is no trace of internal structure. It is, however, frequently the case that a knowledge of the internal structure of a particular plant enables us to interpret certain features in a structureless cast which could not be understood without the help of histological facts. A particularly interesting example of anatomical knowledge affording a key to apparently abnormal peculiarities in a specimen preserved by incrustation, is afforded by the fructification of the genus Sphenophyllum. Some few years ago Williamson described in detail the structure of a fossil strobilus (i.e. cone) from the Coal-Measures, but owing to the isolated occurrence of the specimens he was unable to determine the plant to which the strobilus belonged. On re-examining some strobili of Sphenophyllum, preserved by incrustation, in the light of Williamson’s descriptions, Zeiller was able to explain certain features in his specimens which had hitherto been a puzzle, and he demonstrated that Williamson’s cone was that of a Sphenophyllum. Similar examples might be quoted, but enough has been said to emphasize the importance of dealing as far as possible with both petrifactions and incrustations. The facts derived from a study of a plant in one form of preservation may enable us to interpret or to amplify the data afforded by specimens preserved in another form.

DECORTICATED STEMS.

The fact that plants usually occur in detached fragments, and that they have often been sorted by water, and that portions of the same plant have been embedded in sediment considerable distances apart, is a constant source of difficulty. Deciduous leaves, cones, or angiospermous flowers, and other portions of a plant which become naturally separated from the parent tree, are met with as detached specimens, and it is comparatively seldom that we have the necessary data for reuniting the isolated members. As the result of the partial decay and separation of portions of the same stem or branch, the wood and bark may be separately preserved. Darwin[143] describes how the bark often falls from Eucalyptus trees, and hangs in long shreds, which swing about in the wind, and give to the woods a desolate and untidy appearance. In the passage already quoted from the narrative of the voyage of the Challenger, illustrations are afforded of the manner in which detached portions of plants are likely to be preserved in a fossil state. The epidermal layer of a leaf or the surface tissues of a twig may be detached from the underlying tissues and separately preserved[144]. It is exceedingly common for a stem to be partially decorticated before preservation, and the appearance presented by a cast or impression of the surface of a woody cylinder, and by the same stem with a part or the whole of its cortex intact is strikingly different. The late Prof. Balfour[145] draws attention to this source of error in his text-book of palaeobotany, and gives figures illustrating the different appearance presented by a branch of Araucaria imbricata Pav. when seen with its bark intact and more or less decorticated. Specimens that are now recognised as casts of stems from which the cortex had been more or less completely removed before preservation, were originally described under distinct generic names, such as Bergeria, Knorria and others. These are now known to be imperfect examples of Sigillarian or Lepidodendroid plants. Grand’Eury[146] quotes the bark of Lepidodendron Veltheimianum Presl. as a fossil which has been described under twenty-eight specific names, and placed in several genera.

Since the microscopical examination of fossil plant-anatomy was rendered possible, a more correct interpretation of decorticated and incomplete specimens has been considerably facilitated. The examination of tangential sections taken at different levels in the cortex of such a plant as Lepidodendron brings out the distribution of thin and thick-walled tissue. Regularly placed prominences on such a stem as the Knorria shown in fig. 23 are due to the existence in the original stem of spirally disposed areas of thin-walled and less resistant tissue; as decay proceeded, the thinner cells would be the first to disappear, and depressions would thus be formed in the surrounding thicker walled and stronger tissue. If the stem became embedded in mud or sand before the more resistant tissue had time to decay, but after the removal of the thin-walled cells, the surrounding sediment would fill up the depressions and finally, after the complete decay of the stem, the impression on the mould or on the cast, formed by the filling up of the space left by the stem, would have the form of regularly disposed projections marking the position of the more delicate tissues. The specimen represented in the figure is an exceedingly interesting and well preserved example of a Coal-Measure stem combining in itself representatives of what were formerly spoken of as distinct genera.

Fig. 23. A dichotomously branched Lepidodendroid stem (Knorria mirabilis Ren. and Zeill.). After Renault and Zeiller[147]. (¼ nat. size.) The original specimen is in the Natural History Museum, Paris.

The surface of the fossil as seen at e affords a typical example of the Knorria type of stem; the spirally disposed peg-like projections are the casts of cavities formed by the decay of the delicate cells surrounding each leaf-trace bundle on its way through the cortex of the stem. The surface g exhibits a somewhat different appearance, owing to the fact that we have the cast of the stem taken at a slightly different level. The surface of the thick layer of coal at a shows very clearly the outlines of the leaf-cushions; on the somewhat deeper surfaces b, c and d the leaf-cushions are but faintly indicated, and the long narrow lines on the coal at c represent the leaf-traces in the immediate neighbourhood of the leaf-cushions.

IMPERFECT CASTS.

It is not uncommon among the older plant-bearing rocks to find a piece of sandstone or shale of which the surface exhibits a somewhat irregular reticulate pattern, the long and oval meshes having the form of slightly raised bosses. The size of such a reticulum may vary from one in which the pattern is barely visible to the unaided eye to one with meshes more than an inch in length. The generic name Lyginodendron[148] was proposed several years ago (1843) for a specimen having such a pattern on its surface, but without any clue having been found as to the meaning of the elongated raised areas separated from one another by a narrow groove. At a later date Williamson investigated the anatomy of some petrified fragments of a Carboniferous plant which suggested a possible explanation of the surface features in the structureless specimens. The name Lyginodendron was applied to this newly discovered plant, of which one characteristic was found to be the occurrence of a hypodermal band of strong thick-walled tissue arranged in the form of a network with the meshes occupied by thin-walled parenchyma. If such a stem were undergoing gradual decay, the more delicate tissue of the meshes would be destroyed first and the harder framework left. A cast of such a partially decayed stem would take the form, therefore, of projecting areas, corresponding to the hollowed out areas of decayed tissue, and intervening depressions corresponding to the projecting framework of the more resistant fibrous tissue. A precisely similar arrangement of hypodermal strengthening tissue occurs in various Palaeozoic and other plants, and casts presenting a corresponding appearance cannot be referred with certainty to one special genus; such casts are of no real scientific value[149].

The old generic terms Artisia and Sternbergia illustrate another source of error which can be avoided only by means of a knowledge of internal structure. The former name was proposed by Sternberg and the latter by Artis for precisely similar Carboniferous fossils, having the form of cylindrical bodies marked by numerous transverse annular ridges and grooves. These fossils are now known to be casts of the large discoid pith of the genus Cordaites, an extinct type of Palaeozoic Gymnosperms. Calamites and Tylodendron afford other instances of plants in which the supposed surface characters have been shown to be those of the pith-cast. The former genus is described at length in a later chapter, but the latter may be briefly referred to. A cast, apparently of a stem, from the Permian rocks of Russia was figured in 1870 under the name Tylodendron; the surface being characterised by spirally arranged lozenge-shaped projections, described as leaf-scars. Specimens were eventually discovered in which the supposed stem was shown to be a cast of the large pith of a plant possessing secondary wood very like that of the recent genus Araucaria. The projecting portions, instead of being leaf-cushions, were found to be the casts of depressions in the inner face of the wood where strands of vascular tissue bent outwards on their way to the leaves. If a cast is made of the comparatively large pith of Araucaria imbricata the features of Tylodendron are fairly closely reproduced[150].

A dried Bracken frond lying on the ground in the Autumn presents a very different appearance as regards the form of the ultimate segments of the frond to that of a freshly cut leaf. In the former the edges of the pinnules are strongly recurved, and their shape is considerably altered. Immersed in water for some time fern fronds or other leaves undergo maceration, and the more delicate lamina of the leaf rots away much more rapidly than the scaffolding of veins. Among fossil fern fronds differences in the form of the pinnules and in the shape and extent of the lamina, to which a specific value is assigned, are no doubt in many cases merely the expression either of differences in the state of the leaves at the time of fossilisation or of the different conditions under which they became embedded. Differential decay and disorganisation of plant tissues are factors of considerable importance with regard to the fossilisation of plants. As Lindley[151] and later writers have suggested, the absence or comparative scarcity of certain forms of plants from a particular fossil flora may in some cases be due to their rapid decay and non-preservation as fossils; it does not necessarily mean that such plants were unrepresented in the vegetation of that period. The decayed rhizomes of the Bracken fern often seen hanging from the roadside banks on a heath or moorland, and consisting of flat dark coloured bands of resistant sclerenchyma in a loose sheath of the hard shrivelled tissue, are in striking contrast to the perfect stem. A rotting Palm stem is gradually reduced to a loose stringy mass consisting of vascular strands of which the connecting parenchymatous tissue has been entirely removed. It must frequently have happened that detached vascular bundles or strands and plates of hard strengthening tissue have been preserved as fossils and mistaken for complete portions of plants.

MINERAL DEPOSITS SIMULATING PLANTS.

Apart from the necessity of keeping in view the possible differences in form due to the state of the plant fragments at the time of preservation, and the marked contrast between the same species preserved in different kinds of rock, there are numerous sources of error which belong to an entirely different category. The so-called moss-agates and the well-known dendritic markings of black oxide of manganese, are among the better known instances of purely inorganic structures simulating plant forms.

An interesting example of this striking similarity between a purely mineral deposit and the external form of a plant is afforded by some specimens originally described as impressions of the oldest known fern. The frontispiece to a well-known work on fossil plants, Le monde des plantes avant l’apparition de l’homme[152], represents a fern-like fossil on the surface of a piece of Silurian slate. The supposed plant was named Eopteris Morierei Sap., and it is occasionally referred to as the oldest land plant in books of comparatively recent date. In the Museum of the School of Mines, Berlin, there are some specimens of Angers slate on some of which the cleavage face shows a shallow longitudinal groove bearing on either side somewhat irregularly oblong and oval appendages of which the surface is traversed by fine vein-like markings. A careful examination of the slate reveals the fact that these apparent fern pinnules are merely films of iron pyrites deposited from a solution which was introduced along the rachis-like channel. Many of the extraordinary structures described as plants by Reinsch[153] in his Memoir on the minute structure of coal have been shown to be of purely mineral origin.

The innumerable casts of animal-burrows and trails as well as the casts of egg-cases and various other bodies, which have been described as fossil algae, must be included among the most fruitful sources of error.

It requires but a short experience of microscopical investigation of fossil plant structures to discover numerous pitfalls in the appearance presented by sections of calcareous and siliceous nodules. The juxtaposition of tissues apparently parts of the same plant, and the penetration by growing roots of partially decayed plant débris, serve to mislead an unpractised observer. In sections of the English ‘calcareous nodules’ one very frequently finds the tissue of Stigmarian appendages occupying every conceivable position, and preserved in places admirably calculated to lead to false interpretations. The more minute investigation of tissues is often rendered difficult by deceptive appearances simulating original structures, but which are in reality the result of mineralisation. It is no easy matter in some cases to discover whether a particular cell in a fossil tissue was originally thick-walled, or whether its sclerous appearance is due to the deposition of mineral matter on the inside of the thin cell-membrane. Examples of such sources of error as have been briefly referred to, and others, will be found in various parts of the descriptive portions of this book.

Fig. 24. A. Section of partially disorganised tissue attacked by some boring animal. c, c, coprolites; d, a tunnel made by the borer through the plant tissue.

B. Transverse section of a Lepidodendroid leaf, of which the inner tissues have been destroyed and the cavity filled with coprolites; simulating a sporangium containing spores. (A and B from specimens in the Botanical Laboratory collection, Cambridge.)

TRACES OF WOOD-BORERS IN PETRIFIED TISSUE.

There is one other form of pitfall which should be briefly noticed. In sections of petrified plants one occasionally finds clean cut canals penetrating a mass of tissue, and differing in their manner of occurrence and in their somewhat larger size from ordinary secretory ducts. Such tunnels or canals are probably the work of a wood-boring animal. An example is illustrated in fig. 24 A. Similarly it is not unusual to meet with groups or nests of spherical or elliptical bodies lying among plant tissues, and having the appearance of spores. Such spore-like bodies appear on close examination to be made up of finely comminuted particles of tissue, and in all probability they are the coprolites of some xylophagous animal. Examples of such coprolites are shown in fig. 24 A[154], and in fig. 24 B an interesting manner of occurrence of these misleading bodies is represented. The framework of cells enclosing the nest of coprolites in fig. 24 B, represents the outer tissues of a Lepidodendroid or a Sigillarian leaf; the inner tissues have been destroyed and the cavity is now occupied by what may possibly be the excreta of the wood-eating animal.

Some of the oval spore-like structures met with in plant tissues may, as Renault has suggested, be the eggs of an Arthropod[155]. In a section of a calcareous Coal-Measure nodule in the Williamson collection (British Museum)[156] there occur several fungal spores or possibly oogonia lying among imperfectly preserved Stigmarian appendages. Associated with these are numerous dark coloured and larger bodies consisting of a cavity bounded by a simple membrane; the larger bodies may well be the eggs of some Arthropod or other animal.

PHOTOGRAPHY AND ILLUSTRATION.

In looking through the collections of Coal-Measure plants in the Museums of Berlin, Vienna and other continental towns, one cannot fail to be struck with the larger size of many of the specimens as compared with those usually seen in English Museums. The facilities afforded in the State Collieries of Germany to the scientific investigator may account in part at least for the better specimens which he is able to obtain. It would no doubt be a great gain to our collections of Coal-Measure plants if arrangements could be made in some collieries for the preservation of the finer specimens met with in the working of the seams, instead of breaking up the slabs of shale and consigning everything to the waste heaps. There is one more point which should be alluded to in connection with possible sources of error, and that is the essential importance of accuracy in the illustration of specimens, especially as regard type-specimens. It is often impossible to inspect the original fossils which have served as types, and it is of the utmost importance that the published figures should be as faithful as possible. M. Crépin[157] of Brussels, in an article on the use of photography in illustrating, has given some examples of the confusion and mistakes caused by imperfect drawings. It does not require a long experience of palaeobotanical work to demonstrate the need of care in the execution of drawings for reproduction.