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See also:MAP B .—M,–M... D. G. See also:Distribution of the r:tz alac a E1 ~0 Matnnineae. 1)ible7•irlinae(; D1–D6, Distribution of the Dipteridinae. G5 (urassic ; G12 Cretaceous-See also:Tertiary) ; G1–G17, Distribution of the See also:Ginkgoales G6 (See also:Jurassic and Tertiary) ; G16 (Tertiary, See also:Alaska) ; during the Mesozoic and Tertiary G7 (Jurassic); G14 (Cretaceous-Tertiary) ; Periods. G8 (See also:Rhaetic-Jurassic); G15 (Jurassic) ; G1 (Trias-Tertiary) ; G9 (Trias-Rhaetic) ; G10 (Jurassic, See also:Spitsbergen) ; G2, Ge (Rhaetic-Jurassic); G10 (Rhaetic, See also:Chile); Gu (Jurassic, See also:Franz Josef See also:Land). G4 (Tertiary, See also:Sakhalin I.) ; G11 (Trias) ; as Ctenozamites, Ctenis, and Podozamites, the position of which is ductive See also:organs and stems less certain. Ctenozamites occurs chiefly in the Rhaetic See also:coal-bearing of these fossil Cycads? beds of Scania, and has been found also in the Liassic See also:clays of Cycadean stems have re- cently been found in See also:great abund- ance in Jurassic pants become much more abundant, especially in the See also:Keuper See also:period; as well in See also:North See also:America, from Rhaetic rocks a still greater number of types have been re- throughout See also:Europe. It is corded, among which may be mentioned Nilssonia (fig. To), Anorno- noteworthy that Tertiary zamites, Pterophyllum, Otozamites, Cycadites (fig. II). The See also:species plant-beds have yielded of Nilssonia shown in fig. io (N. compta) is a characteristic member hardly any specimens that of the Jurassic See also:flora, practically identical with a See also:form from Rhaetic can be recognized as rocks described as Nilssonia polymorpha. The large frond of Cycads. Cycadites represented in fig. 11 (C. Saportae) is from the See also:Wealden A more important quesstrata of See also:Sussex, and possibly identical with Cycadites tenuisectus tion is, What knowledge from See also:Portugal. In addition to these genera there are others, such have we of the repro- and possibly higher strata in See also:Wyoming, See also:South Dakota, and other parts of the See also:United States. Cycadean stems have been found also in the upper-most Jurassic, Wealden and See also:Greenland, and other See also:Arctic lands and See also:Dorsetshire and in the Inferior Oolite beds of See also:Yorkshire, as well as in Rhaetic strata in See also:Persia and elsewhere; it is characterized by its bipinnate fronds, and may be compared with the See also:recent Australian genus Bowenia—See also:peculiar among living Cycads in having bipinnate fronds. Ctenis has been incorrectly placed among the ferns by some authors, on See also:account of the occurrence of supposed sporangia on its pinnae; but there is See also:reason to believe that these so-called sporangia are probably nothing more than prominent papillose cells ofythe epidermis. Podozamites (fig. 12) is usually considered to be a Cycad, but the broad pinnae (or leaves) and their arrangement on the See also:axis suggests a possible relationship with the See also:southern coniferous genus Agathis, represented by the Kauri See also:pine and other recent species. The considerable variation in the See also:size of the pinnae of Podozamites, as represented by species from the Jurassic rocks in the Arctic regions and various See also:European localities, recalls the variation in length and breadth of the leaves of Agathis. With regard to the distinguishing features and the distribution of the numerous Cycatean leaves of Mesozoic See also:age, the most striking fact is the abundance of xx. 18 a more compact structure. It is in- FIG. i i.—Cycadites Saporteresting to find that G. R. See also:Wieland of tae: Wealden, See also:England. See also:Lower Cretaceous rocks of England, See also:India and other parts of the See also:world. An example of an See also:Indian Cycadean See also:stem from Upper See also:Gondwana rocks is represented in fig. 13; the See also:surface of the See also:trunk is covered with persistent bases (fig. 13, A) of the fronds known as Ptilophyllum cutchense, which are practically the same as the European species Williamsonia pecten (fig. 17). In a See also:section of the stem (fig. 13, B) a large See also:pith is seen to occupy the axial region, and this is surrounded by a See also:zone of secondary See also:wood, which appears to differ from the characteristic wood of See also:modern Cycads (see See also:GYMNOSPERMS) in having II Yale University has noticed in some of the Cycadean stems from the See also:Black hills of Dakota and Wyoming that the wood appears to possess a similar structure, differing in its narrower medullary rays from the wood of modern Cycads. The See also:lozenge-shaped areas See also:external to the axis of the stem represent the sections of petioles, some of which are shown in fig. 13, A, attached to the stem. The See also:majority of Mesozoic stems agree in external See also:appearance with those of recent species of Encephalartos, Macrozamia, and some other genera; the trunk is encased in a See also:mass of persistent petiole-bases separated from one another by a dense See also:felt or packing of scaly ramenta. The structure of the See also:leaf-stalks is like that of modern
A
B
Cycads, but the ramenta, instead of having the form of See also:long unicellular hairs like those on the petioles and bud-scales of existing species are exactly like the paleae or ramental scales characteristic of the majority of ferns. This See also:fern-like See also:character affords an interesting survival of the See also:close relationship between Cycads and Ferns. Some examples of Jurassic Cycadean stems from Wyoming are characterized by an unusually See also:rich development of ramental scales; the ramenta from the old leaf-bases form an almost See also:complete covering over the surface of the -trunk. See also:Professor Lester See also: The best preserved specimens of the true Bennettites type so far described are from the Lower See also:Green-See also:sand and Wealden of England, and from Upper Mesozoic strata in North America, See also:Italy and See also:France. A study of the anatomical structure of the FIG. 1 — Cycadeoidea vegetative stem, which on the whole eigantea. Portland rocks, is very similar to that 9f recent England. Cycads (fig. 15, i and 2), reveals certain characters which are not met with in modern Cycads. The See also:chief distinguishing feature is afforded by the leaf-traces; in recent species (ssee GYMNOSPERMS)these pursue a somewhat complicated course as they pass from the petiole towards the vascular See also:cylinder of the stem, but in Bennettites the vascular bundles from the leaves followed a more See also:direct course through the cortex of the stem (fig. 15, 3). Among existing types the genus Macrozamia appears to show the nearest approach to this simpler structure of the leaf-traces. In a Floridan species of Zamia the leaf-traces are described as characterized by a more direct course from the See also:stele of the stem to the leaves than in most modern genera, thus agreeing more closely with the See also:extinct Bennettites. The typical Bennettites See also:female flower (fig. 15, 4 and 7), as investigated in See also:English. See also:French, See also:Italian, and See also:American specimens, may be briefly described as a See also:short lateral shoot or peduncle, arising in a leaf-axil and terminating in a bluntly rounded See also:apex, bearing numerous linear bracts enclosing a central group of appendages, some of which consist of slender pedicels traversed by a vascular strand and bearing a single terminal ovule enclosed in an integument, which forms a distal See also:canal or micropyle. Associated with these seminiferous pedicels occur sterile appendages consisting of slender stalks, terminating in distal expansions, which form a fleshy covering over the surface of the flower, leaving small apertures immediately above the micropyles for the entrance of the See also:pollen-grains. It has been suggested by some authors that the almost complete investment of the small Bennettites seeds by the surrounding swollen ends of the interseminal scales (fig. 15, 7) represents an approach to the angiospermous ovary. In Bennettites the ovules are See also:left exposed at the apex, but they are by no means so distinctly gymnospermous as in recent Cycads and Conifers. The seeds have in some cases been preserved in wonderful perfection, enabling one to make out the structure of the embryo, with its bluntly conical radicle and two fleshy cotyledons filling the exalbuminous See also:seed (fig. 1 II). Our knowledge of the reproductive organs of the Bennettitaceae has until recently been confined to the female flowers, as described by Carruthers, Solms-Laubach, Lignier, and others. The fortunate See also:discovery of several See also:hundred Cycadean stems in the United States, of Lower Cretaceous and Upper Jurassic age, has supplied abundant material which has lately been investigated and is still receiving See also:attention at the hands of Mr Wieland. This investigator has already published a well-illustrated account of his discoveries, which give valuable See also:information as to the See also:morphology of the male organs, and See also:lead us to expect additional results in the future of the greatest importance and See also:interest. On some of the American stems flowers have been found, See also:borne at the apex of lateral shoots, which possess fully See also:developed male organs consisting of sporangia with spores (pollen-grains), surrounding a conical central receptacle bearing numerous small and probably functionless or immature ovules (fig. 15, lo). The structure of this type of flower may be briefly described as follows. In shape and size the flower is similar to that long known as the female flower of Bennettites and Williamsonia. A number of hairy linear bracts enclose the whole; See also:internal to these occur 12 to 20 crowded pinnate leaves (sporophylls), with their apical portions See also:bent over towards the axis of the flower, the bases of the petioles being fused laterally into a disk surrounding the See also:base of the conical receptacle. Numerous pairs of pinnules are attached to the rachis of each sporophyll, and the larger pinnules See also:bear 20 to 30 synangia (sori or plurilocular sporangia) (fig. 15, 8 and 9). The synangia consist of a stout See also:wall composed of thick-walled cells, succeeded by a layer of more delicate and smaller elements; and internal to the wall occur two rows of sporangial loculi containing microspores. When the synangia are ripe dehiscence takes See also:place along a median See also:line between the two rows of loculi. In size, position, arrangement, and manner of dehiscence the sporangia bear a striking resemblance to those of Marattia and Danaea among recent Marattiaceae. The most important point elucidated by this discovery is the very close See also:correspondence of the male organs of the Bennettites flower with the sporophylls and synangia of Marattiaceous ferns—a further relic of the See also:common origin of Cycads and Ferns. It remains to be seen if the ovuliferous See also:cone in the centre of the flower represents simply a functionless gynoecium, as in Welwitschia and abnormal cones of certain Coniferae, or if the flowers were hermaphrodite, with both male and female organs fully developed. We have a See also:combination in the same flower of stalked ovules, the structure of which has already been described, and interseminal scales constituting a complex gynoecium, which exhibits in certain features an approach to the angiospermous type, and differs in structure from other Gymnosperm flowers, associated with male organs constructed on a See also:plan almost identical with that of the sporophylls in Marattiaceae. In many of the flowers de-scribed by Mr Wieland the structure is identical in essential features with that of the female flowers of Bennettites Gibsonianus described by Carruthers and by Solms-Laubach, and with that of a French Liassic species described by Lignier: the whole consists of a See also:convex receptacle bearing mature seeds at the tips of pedicels associated with interseminal scales (fig. 15, 7) as already described. Mr Wieland's researches have, however, demonstrated the existence in flowers of this type of the remains of a disk at the base of the receptacle, between the receptacle and the surrounding bracts, to which staminate leaves were originally attached. The flowers hither-to regarded as female were in some cases at least hermaphrodite, but the male organs had been thrown off before the complete development of the gynoecium. This fact suggests the possibility that the flowers described by Mr Wieland, in which the male organs are mature and the gynoecium is composed of very short and immature ovuliferous stalks and interseminal scales, are not essentially distinct from those which have lost the staminate leaves I, Bennettites stem: portion of transverse section of stem; a, vascular cylinder; b, leaf-traces; c, pith; d, cortex. 2, Bennettites stem, tangential section; e, flower-peduncles. 3, Bennettites stem, leaf-traces attached to the vascular cylinder and passing as See also:simple strands through the cortex; d, cortex. 4, Williamsonia, Wealden, England. 5, See also:Young leaf of Bennettites. 6, Ramenta of Bennettites in transverse section. 7, Bennettites, female flower in See also:longitudinal section; f, apex of peduncle; g, bracts (shown in surface view in 4); h, seeds and seminiferous pedicels; i, interseminal scales. 8, Bennettites, synangium of male flower, showing line of dehiscence, k, and microspores, 1. 9, Synangium, in transverse section, showing sporangial See also:groups, m, and microspores, 1. I0, Bennettites flower in See also:vertical section, showing the central female portion, n, two sporophylls bearing synangia (male), o, and hairy bracts, g. II, Bennettites seed in longitudinal section, showing the dicotyledonous embryo; p, cotyledons; r, radicle; s, testa. (1-3, after Carruthers; 5, 8, 9 and to, after Wieland; 7, after See also:Scott; I1, after Solms-Laubach.) and possess mature seeds. It is probable that the flowers of Bennettites were normally hermaphrodite, and they may have been markedly protandrous. We cannot decide at See also:present whether the gynoecium in a flower, such as that represented in fig. 15, 7, has partially aborted or whether it would have matured later after the fall of the male organs. It is clear that Bennettites differed in many essential respects from the few modern survivors of the See also:Cycadophyta. Fossil flowers of a type more like that of modern Cycads are few in number, and it is not by any means certain that all of those described as Cycadean flowers and seeds were borne by plants which should be included in the Cycadophyta; a few female flowers have been described from Rhaetic rocks of Scania and elsewhere under the name Zamiostrobus—these consist of an axis with slender pedicels or carpophylls given off at a wide See also:angle and bearing two ovules at the distal end ; the structure is in fact similar to that of a Zamia female flower, in which the internodes of the peduncle have been elongated so as to give a looser arrangement to the carpels. It has been suggestedthat one at least of the flowers, that originally described by Mr Carruthers from the Inferior Oolite of Yorkshire as Beania gracilis, may have been borne by a member of the Ginkgoales. From Jurassic rocks of France and Italy a few imperfect specimens have been described as carpels of Cycads, like those of the recent genus Cycas (see GYMNOSPERMS); while a few of these may have been correctly identified, an inspection of some of the See also:original examples in the See also:Paris collections leads one to See also:express the See also:opinion that others are too imperfect to determine. Pinnate fronds of the Cycas type, characterized by the presence of a midrib and no lateral See also:veins in the linear pinnae, are recorded from Rhaetic rocks of See also:Germany, from Wealden strata in England (fig. II) and Portugal, and from Liassic beds in Dorsetshire. One large specimen is figured by Heer from Lower Cretaceous rocks of Greenland, and by the See also:side of the frond is shown a carpel with lateral ovules, as in the female flower of Cycas; but an examination of the type-specimen in the See also:Copenhagen Museum led the present writer to regard this supposed carpel as valueless. Professor Nathorst, as the result of a more recent examination of Heer's specimen, found that the segments of the frond are characterized by the presence of two parallel veins instead of a single midrib, with a See also:row of stomata between them; for this type of Cycadean leaf he proposed the generic name Pseudocycas. Another well-known Cycadean genus is Williamsonia, so named by Mr Carruthers in 187o, and now applied to certain pinnate fronds — e.g. those pre- viously described as Zamites gigas (fig. 16), and others known under such names as Pterophyllum or Ptilophyllum pecten, &c., both common Jurassic species — as well as to stems bearing peduncles with terminal See also:oval flowers, similar in form to those of Bennettites. There is See also:good See also:evidence for supporting Professor See also:Williamson's conclusions as to the organic connexion between the flowers, originally de-scribed from Inferior Oolite rocks of Yorkshire and subsequently named Williamsonia (fig. 15, 4), and the fronds of Zamites gigas, now known as Williamsonia gigas (fig. 16). There can be little doubt that the majority of the Cycadean fronds of Jurassic and Wealden age, which are nearly always found detached from the See also:rest of the plant, were borne on stems of the Bennettites type. Williamson was the first to express the opinion that the Bennettitean flowers known as Williamsonia were borne on the trunks which terminated in a See also:crown of pinnate fronds FIG. 16.—Frond of Williamsonia gigas. of the type long known as Inferior Oolite, England. Zamites gigas; this view was regarded by Saporta and others as incorrect, and the nature of the Bennettitean foliage was left an open question. A re-examination of the English material in the museums of Paris and else-where has confirmed Williamson's conclusions. Mr Wieland has also described young bipinnate fronds, very like those of recent species of Zamia and Encephalartos, attached to a Bennettites stem, and exhibiting the vernation characters of many recent Cycads (fig. 15, 5). In Williamsonia the stem See also:bore comparatively long fertile shoots, which, in contrast to those of Bennettites, projected several inches beyond the surface of the See also:main trunk, and terminated in a flower which appears to have resembled those of the true Bennettites. Nathorst has recently described specimens of Williamsonia from the Jurassic rocks of See also:Whitby with micro-Sporophylls like those of Wieland's species. Williamsonia occurs in the Upper Gondwana rocks of India; it is recorded also from strata ranging from the Rhaetic to the Lower Cretaceous period in England, Portugal, See also:Sweden, See also:Bornholm, Greenland, Italy and North America. Professor Nathorst has described another type of stem from the Rhaetic beds of Scania. It consists of a comparatively small and repeatedly forked axis bearing in each See also:fork a flower; the flowers, which are regarded as male and female, appear to be similar to those of Bennettites. The leaves, borne on the regions between the false dichotomies, are those of Anomozaanites See also:minor, a type of Cycadean frond originally determined by See also:Brongniart. The flowers, or some of them, were originally described by Nathorst as Williamsonia angustifolia. This form of stem, of a See also:habit entirely different from that of recent Cycads and extinct Bennettites, points to the existence in the Mesozoic era of , in addition to several well-preserved female flowers. C. A. Hollick another type of Gymnosperm allied to the Bennettitales of the Jurassic and Cretaceous periods by its flowers, but possessing a distinctive character in its vegetative organs. There is no doubt that the Cycadophyta, using the See also:term suggested by Nathorst in 1902, was represented in the Mesozoic period by several distinct families or classes which played a dominant See also:part in the floras of the world before the See also:advent of the See also:Angiosperms. In addition to the bisporangiate reproductive shoots of Bennettites, distinguished by many important features from the flowers of recent Cycads, a few specimens of flowers have been discovered exhibiting a much closer resemblance to those of existing Cycads, e.g. Androstrobus Balduini from Bathonian rocks of France; Zamites familiaris, described many years ago by Corda, from Lower Cretaceous rocks of Bohemia, and Androstrobus Nathorsti, from Wealden beds in Sussex. The majority of the species were, however, characterized by flowers of a different type known as Bennettites and Williamsonia. The living See also:Maidenhair-See also:tree (Ginkgo biloba) (see GvMNOSPERMS) remains, like Matonia and Dipteris, among the ferns, as an isolated relic in the midst ainkgoales. of recent vegetation. In Rhaetic, Jurassic and Wealden floras, the Ginkgoales were FIG. 18.—Leaves of Ginkgoales. exceedingly abundant (Map B, GI–Gl,) ; in addition to leaves agreeing almost exactly with those of the recent A, Ginkgodiunx, See also:Japan (Jurassic). species (fig. 18), there are others separated as a distinct B, C, D, E, F, H, Ginkgo leaves.—B, from Franz Josef Land (Jurassic); genus, Baiera (fig. 18, G), characterized by the greater C, Greenland (Lower Cretaceous) ; D, See also:Siberia (Jurassic) ; E, Germany number and narrower form of the segments, which may (Wealden); F, England (Jurassic); H, See also:China (Rhaetic). be best compared with such leaves as those of the G, Baiera leaf, Inferior Oolite, England. recent fern Actiniopteris and of certain species of Schizaea. (A, after Yokoyama ; B, after Nalhorst; C, D, after Heer; E. after Schenk ; Male flowers, like those of Ginkgo biloba, but usually H, after Krasser. All the figures z nat. size.) characterized by a rather larger number of oval pollen-sacs on the stamens, have been found in England, Germany, Siberia and elsewhere in association with Ginkgo and Baiera foliage. The occasional occurrence of three or even four pollen-sacs on the stamens of the recent species affords a still closer agreement between the extinct and living types. Seeds like those of Ginkgo biloba have also been recorded as fossils in Jurassic rocks, and it is possible that the type of flower known as Beania, from the Inferior Oolite rocks of Yorkshire, may have been borne by Ginkgo or Baiera. The regions from which satisfactory examples of Ginkgoales (Baiera or Ginkgo) have been recorded are shown in Map B (G1-G,). Both Tertiary and Mesozoic localities are indicated in the map. An adequate account of fossil Mesozoic Conifers is impossible within the limits of this See also:article. Coniferous twigs are very common in Mesozoic strata, but in most cases we are compelled Coniferales. to refer them to provisional genera as the evidence of vegetative shoots alone is not sufficient to enable us to determine their position within the Coniferae. There are, however, several forms which it is reasonable to include in the Araucarieae; that this See also:family was to the fore in the vegetation of the Jurassic period is unquestionable. We have not merely the striking resemblance of vegetative shoots to those of recent species of See also:Araucaria and Agathis, e. . species of Nageiopsis, abundantly represented in the Upper Jurassic beds of the See also:Potomac See also:area in North America, species of Pagiophyllum and other genera of Jurassicand Wealden age, but an abundance of fossil wood (Araucarioxylon) from Jurassic and Cretaceous strata in Europe, North America, See also:Madagascar and elsewhere agreeing with that of recent Araucarieae, and E. C. See also:Jeffrey have recently shown that some Lower Cretaceous specimens of the well-known genus Brachyphyllum obtained from Staten See also:Island, N.Y., possess wood of the Araucarian type. This genus has long been known as a common and widely spread Jurassic and Cretaceous conifer, but owing to the See also:absence of petrified specimens and of well-preserved cones, it has been impossible to refer it to a definite position in the Coniferales. It is now clear that some at least of the species of Brachyphyllum must be referred to the Araucarieae. In a recently published See also:paper See also:Seward and See also:Ford have given a See also:general account of the Araucarieae, recent and extinct, to which reference may be made for further details as to the See also:geological See also:history of this See also:ancient section of the Coniferales. Some of the fossils referred to the genus Kaidocarpon, and originally described as monocotyledonous inflorescences, are undoubted Araucarian cones; other cones of the same type have been placed in the genus Cycadeostrobus and referred to Cycads. Araucarites Hudlestoni, described by Mr Carruthers from the Coralline Oolite rocks of See also:Malton in Yorkshire; Araucarites sphaerocarpa from the Inferior Oolite of See also:Somerset; also another cone found in the See also:Northampton Sands, which is probably specifically identical with A. Hudlestoni, and named by Carruthers Kaidocarpon ooliticum, afford good illustrations of See also:British Araucarian flowers. A flower of a rather different type, Pseudaraucaria See also:major, exhibiting in the occurrence of two seeds in each See also:scale an approach to the cones of Abietineae, has been described by Professor Fliche from Lower Cretaceous rocks of See also:Argonne. The well-known Whitby See also:jet of Upper Liassic age appears to have been formed to a large extent from Araucarian wood. Among the more abundant Conifers of Jurassic age may be mentioned such genera as Thuytes and Cupressites, which agree in their vegetative characters with members of the Cupressineae, but our knowledge of the cones is far from satisfactory. Many of the small female flowers borne on shoots with foliage of the Cupressus type consist of spirally disposed and not verticillate scales, e.g. Thuytes expansus, a common Jurassic species. Fossil wood, described under the name Cupressinoxylon, has been recorded from several Mesozoic horizons in Europe and elsewhere, but this term has been employed in a wide sense as a designation for a type of structure met with not only in the Cupressineae, but in members of other families of Coniferae. The Abietineae do not appear to have played a prominent part before the Wealden period; various older species, e.g. Rhaetic specimens from Scania, are recorded, but it is not until we come to the Upper Jurassic and Wealden periods that this modern family was abundantly represented. Fossil wood of the Pinites type (Pityoxylon) has been described from England, France, Germany, Sweden, Spitsbergen, North America and elsewhere; some of the best British examples have been obtained from the so-called Pine-raft, the remains of See also:water-logged and petrified wood of Lower See also:Greensand age, seen at See also:low water near See also:Brook Point in the Isle of See also:Wight. Well-preserved Abietineous female flowers have been obtained from the Wealden rocks of England and See also:Belgium, e.g. Pinites. Dunkeri, P. Solmsi, &c.; specimens of seeds and vegetative shoots are recorded also from Spitsbergen and other regions. Hollick and Jeffrey have recently added to our knowledge of the See also:anatomy of Cretaceous species of Pinus, and See also:Miss Stopes and Dr Fujii have made imortant contributions on the structure of Cretaceous plants from Japan. Cones of Lower Cretaceous age have been described by Fliche from Argonne, which bear a close resemblance to the female flowers of recent species of Cedrus. The two surviving species of See also:Sequoia afford an See also:illustration of the persistence of an old type, but unfortunately most of the Mesozoic species referred to this genus do not possess sufficiently perfect cones to confirm their See also:identification as examples of Sequoia. Some of the best examples of cones and twigs referred to Sequoia are those described by Heer from Cretaceous rocks of Greenland, and Professor D. P. See also:Penhallow of See also:Montreal has described the anatomical structure of the stem of Sequoia Langsdorfii, a Tertiary species occurring in Europe and North America. There are a few points suggested by a general survey of the Mesozoic floras, which may be briefly touched on in conclusion. In following the progress of plant-See also:life through those periods in the history of the See also:earth of which records are left in ancient sediments, seams of coal or old land-surfaces, we recognize at certain stages a want of continuity between the floras of successive ages. The imperfection of the geological See also:record, considered from the point of view of See also:evolution, has been rendered See also:familiar by See also:Darwin's remarkable See also:chapter in the Origin of Species. Breaks in the See also:chain of life, as represented by gaps in the blurred and incomplete documents afforded by fragmentary fossils, are a necessary consequence of the general plan of geological evolution; they See also:mark missing chapters rather than sudden breaks in an evolutionary See also:series. On the other See also:hand, a study of the plant-life of past ages tends to the conviction that too much stress may be laid on the imperfection of the geological record as a See also:factor in the See also:interpretation of palaeontological data. The See also:doctrine of Uniformitarianism, as propounded by See also:Lyell, served to establish See also:geology on a firmer and more rational basis than it had previously possessed; but latterly the tendency has been to modify the Lyellian view by an See also:admission of the See also:probability of a more intense See also:action of groups of forces at certain stages of the earth's history. As a definite instance a short See also:review may be given of the evidence of palaeobotanical records as regards their bearing on plant-evolution. Starting with the Permo-Carboniferous vegetation, and omitting for the moment the Glossopteris flora, we find a comparatively homogeneous flora of wide See also:geographical range, consisting to a large extent of arborescent lycopods, calamites, and other vascular cryptogams, plants which occupied a place comparable with that of Gymnosperms and Angiosperms in our modern forests; with these were other types of the greatest phylogenetic importance, which serve as See also:finger-posts pointing to lines of evolution of which we have but the faintest signs among existing plants. Other types, again, which may be referred to the Gymnosperms, played a not unimportant part in the Palaeozoic vegetation. No conclusive See also:proof has so far been adduced of the existence in those days of the Cycads, nor is there more than partial evidence of the occurrence of genera which can be placed with confidence in any of the existing families of Conifers. There are, moreover, no facts furnished by fossil plants in support of the view that Angiosperms were represented either in the low-lying forests or on the slopes of the mountains of the Coal period. Passing higher up the geological series, we find but scanty records of the vegetation that existed during the closing ages of the See also:Permian period, and of the plants which witnessed the beginning of the Triassic period we have to be content with the most fragmentary See also:relics. It is in rocks of Upper Triassic and Rhaetic age that abundant remains of rich floras are met with, and an examination of the general features of the vegetation reveals a striking contrast between the Lower Mesozoic plants and those of the Palaeozoic period. Arborescent Pteridophytes are barely represented, and such dominant types as Lepidodendron, Sigillaria, Calamites and Sphenophyllum have practically ceased to exist; Cycads and Conifers have assumed the leading role, and the still luxuriant fern vegetation has put on a different aspect. This description applies almost equally to the floras of the succeeding Jurassic and Wealden periods. The See also:change to this newer type of vegetation was no doubt less sudden than it appears as read from palaeobotanical records, but the transition period between the Palaeozoic type of vegetation and that which flourished in the Lower Mesozoicera, and continued to the close of the Wealden age, was probably characterized by rapid or almost sudden changes. In the southern hemisphere the Glossopteris flora succeeded a Lower Carboniferous vegetation with a rapidity similar to that which marked the passage in the north from Palaeozoic to Mesozoic floras. This apparently rapid alteration ih the character of the southern vegetation took place at an earlier period than that which witnessed the transformation in the See also:northern hemisphere. The appearance of a new type of vegetation in India and the southern hemisphere was probably connected with a widespread lowering of temperature, to which reference has already been made. It was from this Glossopteris flora that several types gradually migrated across the See also:equator, where they formed part of the vegetation of more northern regions. The difference between the Glossopteris flora and those which have left traces in the Upper Gondwana rocks of India, in the Wianamatta and Hawkesbury beds of See also:Australia, and in the Stormberg series of South See also:Africa is much less marked than that between the Permo-Carboniferous flora of the northern hemisphere and the succeeding Mesozoic vegetation. In other words, the change took place at an earlier period in the south than in the north. To return to the northern hemisphere, it is clear that the Wealden flora, as represented by plants recorded from England, France, Belgium, Portugal, See also:Russia, Germany and other European regions, as also from Japan and elsewhere, carries on, with minor See also:differences, the facies of the older Jurassic floras. It was at the close of the Wealden period that a second evolutionary See also:wave swept over the vegetation of the world. This change is most strikingly illustrated by the inrush of Angiosperms, in the equally marked decrease in the Cycads, and in the altered character of the ferns. It would appear that in this See also:case the new See also:influence, supplied by the advent of Angiosperms, had its origin in the north. Unfortunately, our knowledge of the later floras in the southern hemisphere is very incomplete, but a similar transformation appears to have characterized the vegetation south of the equator. As to the nature of the chief factors concerned in the two revolutions in the See also:vegetable See also:kingdom, if it is admissible to use so strong a term, only a guess can be hazarded. See also:Physical conditions no doubt played an important part, but whatever cause may have had the greatest See also:share in disturbing the See also:equilibrium of evolutionary forces, it would seem that the apparently sudden appearance of Cycads and other types at the close of the Palaeozoic period made a widespread and sudden impression on the whole character of the vegetation. At a later See also:stage—in See also:post-Wealden days—it was the appearance of Angiosperms, probably in northern latitudes, that formed the chief See also:motive See also:power in accelerating the transition in the facies of plant-life from that which marked what we have called the Mesozoic floras, to the vegetation of the Upper Cretaceous and Tertiary periods. With the advent of Angiosperms began, as the See also:late See also:marquis of Saporta expressed it, " Une revolution, ainsi rapide dans sa See also:marche qu'universelle dans ses effets." From the floras of the Tertiary age we pass by See also:gradual stages to those which characterize the present phase of evolutionary progress. Among modern floras we find here and there isolated types, such as Ginkgo, Sequoia, Matonia, Dipteris and the Cycads, persisting as more successful survivals which have held their own through the course of ages; these plants remain as vestiges from a remote past, and as links connecting the vegetation of to-See also:day with that of the Mesozoic era. AuTHoruTlEs.—Glossopteris Flora: See also:Blanford, H. F., " On the age and correlation of the Plant-bearing Series of India, &c.," Quarterly See also:Journal Geol. See also:Soc. xxxi. (1875); Feistmantel, " Fossil Flora of the Gondwana See also:System," Mem. Geol. Sure. India, vols. iii., &c. (1899, &c.) ; Seward, Fossil Plants as Tests of See also:Climate (See also:Cambridge, 1892), with bibliography; " The Glossopteris Flora," See also:Science Progress, with bibliography; " On the Association of Sigillaria and Glossopteris in South Africa," Q.J.G.S., vol. H. (1897); E. A. N. See also:Arber, See also:Catalogue of the Fossil Plants of the Glossopteris Flora in the See also:Department of Geology (British Museum, Nat. Hist., Brit. See also:Mus. Catalogue (See also:London, r9o5), with full bibliography; Medlicott and Blanford, See also:Manual of the Geology of India (2nd ed., See also:Oldham, R. D., See also:Calcutta, 1893) ; See also:David, " Evidences of Glacial Action in Australia in Permo-Carboniferous See also:time," Q.J.G.S., vol. Iii. (1896); Zeiller, Elements de paleubotanique (Paris, 19:>0) ; Potonie, " Fossile Pflanzen 550 aus See also:deutsch and portugiesisch Ostafrika," Deutsch-Ostafrika, vii. (See also:Berlin, 1900), with bibliography. General : Potonie, Lehrbuch der Pflanzenpalaeontologie (Berlin, 1899) ; Scott, Studies in Fossil See also:Botany (1900) ; Seward, Fossil Plants (Cambridge: vol. i., 1898) ; vol. ii. 1910, with bibliography; Zeiller, " Revue See also:des travaux de paleontologie vegetate," Rev. gen. bot. (1903) et seq. Catalogue of the Mesozoic Plants in the British Museum, (a) " Wealden Flora," pts. i. and ii.; (b) " Jurassic Flora," pt. i. (1894–1901), pt. ii. (1904), with bibliography; " On the Structure and See also:Affinities of Matonia pectinata, with Notes on the Geological History of the Matonineae," Phil. Trans. cxci. (1899) ; " On the Structure. &c., of Dipteris," ibid. cxciv. (1901, with bibliography; Seward and Ford, " The Araucarieae, recent and extinct," Phil. Trans. R. Soc. (London, 1906) ; G. R. Wieland, "American Fossil Cycads," Publication See also:Carnegie Instil. (See also:Washington, 1906) ; Nathorst, " Palaobotanische Mittel," K. Svensk. Vetenskaps. Akad. Hand. xlii., No. 5 (1907) ; The See also:Norwegian North-Polar Expedition, iii. (1893–1896); " Fossil Plants from Franz Josef Land;" L. F. Ward, " Status of the Mesozoic Floras of the United States," Twentieth See also:Ann. See also:Rep. Geol. Survey (Washington, 1900) ; Solms-Laubach, " Ueber das Genus Pleuromeia," Bot. Zeit. (1899) ; See also:Newton and T'eall, " Notes on a Collection of Rocks and Fossils from Franz Josef Land," Q.J.G.S. liii. (1897) ; Hollick and Jeffrey, " Studies of Cretaceous Coniferous remains, Mem. New See also:York Botanical See also:Garden, vol. iii. (1909); Stopes and Fujii, " Structure and Affinities of Cretaceous Plants," Phil. Trans. R. Soc. (1910). References to important papers on Mesozoic botany will be found in the See also:bibliographies mentioned in the above See also:list. (A. C. SE.) After the Wealden period, and before the deposition of the lowest strata of the See also:Chalk, so remarkable a change takes place in the character of the vegetation that this break Lower must be taken as, botanically, the transition point Cretaceous. from a Secondary to a Tertiary flora. A flora consisting entirely, with a single doubtful exception, of Gymnosperms and Cryptogams gives place to one containing many flowering plants; and these increase so rapidly that before long they seem to have crowded out many of the earlier types, and to have themselves become the dominant forms. Not only do Angiosperms suddenly become dominant in all known plant-bearing deposits of Upper Cretaceous age, but strangely enough the earliest found seem to belong to living orders, and commonly have been referred to existing genera. From Cretaceous times onwards See also:local distribution may change; yet the successive floras can be analysed in the same way as, and compared with, the living floras of different regions. World-wide floras, such as seem to characterize some of the older periods, have ceased to be, and plants are distributed more markedly according to geographical provinces and in See also:climatic zones. This being the case, it will be most convenient to discuss the Tertiary floras in successive See also:order of appearance, since the main interest no longer lies in the occurrence of See also:strange extinct plants or of transitional forms connecting orders now completely isolated. The accurate correlation in time of the various scattered plant-bearing deposits is a See also:matter of considerable difficulty, for plant-remains are preserved principally in lacustrine strata laid down in separate basins of small' extent. This it is obvious must commonly be the case, as most leaves and fruits are not calculated to See also:drift far in the See also:sea without injury or in abundance; nor are they likely as a rule to be associated with marine organisms. Deposits containing marine fossils can be compared even when widely separated, for the ocean is continuous and many marine species are world-wide. Plants, on the other hand, like land and fresh-water animals, occupied areas which may or may not have been continuous. Therefore, without a knowledge of the physical See also:geography of any particular period, we cannot know whether like or unlike floras might be expected in neighbouring areas during that period. If, however, we discover plant-bearing strata interstratified with deposits containing marine fossils, we can See also:fix the period to which the plants belong, and may be able to correlate them in distinct areas, even though the floras be unlike. This clear stratigraphical evidence is, however, so rarely found that much uncertainty still remains as to the true age of several of the floras now to be described. In rocks approximately See also:equivalent to the Lower Greensand of England, or slightly earlier, Angiosperms make their first appearance; but as the only strata of this age in See also:Britain are of marine origin, we have to turn to other countries for the evidence. [TERTIARY The earliest Angiosperm yet found in Europe is a single mono- cotyledonous leaf of doubtful affinities, named by Saporta Alismacites primaevus (fig. I), and found in the Valenginian strata of Portugal. These deposits seem to be equivalent to British Wealden rocks, though in the latter, even in their upper part, no trace of Angiosperms has been discovered. No other undoubted Angiosperm has yet been discovered in Europe in strata of this age, but Heer records a See also:poplar-like leaf from Urgonian strata, a stage newer than the Valenginian, in Greenland, and Saporta has described from strata of the same date in Portugal a Euphorbiaceous plant apparently closely allied to the living Phyllanthus and named by him Choffatia Francheti (fig. 2). We FIG. 1. Alismacites must turn to North America for a See also:fuller primaevus. knowledge of the earlist flowering-plants. In S. Dakota a remarkable series has been discovered, lying unmistakably between marine Upper Jurassic rocks below and Upper Cretaceous above. There has been a certain amount of confusion as to the exact strata in which CreAmerican taceous. the plants occur, but this has now been cleared up by the researches of Lester F. Ward, who has shown how the Secondary flora gives place to one of Tertiary character. The lower strata—i.e. those most allied to the Jurassic—contain only Gymnosperms and Cryptogams. The next See also:division (Dakota fined mainly to the Older Potomac, FIG. 2.—Choffatia Francheti. while the See also:Dicotyledons are principally represented in the Newer Potomac, though occurring more rarely even down to the base of the series. Six successive stages have been defined in the Potomac formation. The See also:Mount See also:Vernon beds, which occur about the See also:middle of the series, have as yet yielded only a small number of species, though these include the most interesting See also:early Angiosperms. Among them are recorded a See also:Casuarina, a leaf of Sagittaria (which however, as observed by Zeiller, may belong to Smilax), two species of poplar-like leaves with remarkably cordate bases, Menispermites (possibly a water-See also:lily) and Celastrophyllum (perhaps allied to Celastrus). Proteophyllum, found in the same See also:bed, and also in the Infra-Cretaceous of Portugal, seems to have belonged to a Proteaceous plant, though only leaves without fruits have yet been discovered in deposits of this early date. Whatever doubt may be left as to the exact botanical position of these early Lower Cretaceous Angiosperms, it is clear that both Monocotyledons and Dicotyledons are represented by several types of leaves, and that the flora ex-tended over wide areas in North America and Greenland, and is found again at a few points in Europe. There is yet no clear evidence either of climatic zones or of the existence of geographical provinces during this period. The next strata, the See also:Aquila See also:Creek series, contain a well-marked dicotyledonous flora, in which both the form and nervation of the leaves begin to approximate to those of recent times. The leading' characteristic of this Middle Potomac flora is the proportion of Dicotyledons. Notwithstanding this apparent passage-bed, there is a marked difference between the Older and the Newer Potomac floras, very few species passing from the one to the other. Only 15 out of 405 plants in the older series occur in the beds above No. 2 of See also:Meek and See also:Hayden) contains Gymnosperms and Ferns of Neocomian types, or even of Neocomian species; but mingled with these occur a few dicotyledonous leaves belonging to four genera. The specimens are very fragmentary, and all that can be said is that one of the forms may be allied to See also:oak, another to fig, a third to Sapindus, and the See also:fourth may perhaps be near to See also:elm. The " Potomac Formation " of See also:Virginia and See also:Maryland is doubtless also mainly of Neocomian age, for though it rests unconformably on much older strata, the successive floras found in it are so allied to those of S. Dakota as to leave little doubt as to the general homotaxis of the series. Lester Ward re-cords no fewer than 737 distinct forms, consisting chiefly of Ferns, Cycads, Conifers and Dicotyledons, t r and cads bei See also:con-e s g though already more than 35o species have been determined from this newer series. The plants from the Amboy Clays, which form the most important division of the Newer Potomac series and were monographed in 1895 by J. S. See also:Newberry, seem to belong to the commencement of the Upper Cretaceous period. It is remarkable that nearly 8o% of the species are Dicotyledons, and that no Monocotyledons have been found. The See also:mere enumeration of the genera will indicate how close the flowering plants are to living forms. Newberry records Juglans, Myrica (7 species), Populus, Salix (5 species), Quercus, Planera, Ficus (3 species), Persoonia and another extinct Proteaceous genus named Proteoides, See also:Magnolia (7 species), Liriodendron (4 species), Menispermites, Laurus and allied plants, Sassafras (3 species), Cinnamomum, Prunus, Hymenaea, Dalbergia, Bauhinia, Caesalpinia, Fontainea, Colutea and other See also:Leguminosae, Ilex, Celastrus, Celastrophyllum (to species), Acer, Rhamnites, Paliurus, Cissites, Tiliaephyllum, Passiflora, See also:Eucalyptus (5 species), Hedera, Aralia (8 species), Cornophyllum, See also:Andromeda (4 species), Myrsine, Sapotacites, Diospyros, Acerates, See also:Viburnum and various genera of uncertain affinities. The points that suggest themselves with regard to this flora are, that it includes a See also:fair See also:representation of the existing orders of warm-temperate See also:deciduous trees; that the more See also:primitive types—such as the Amentaceae—do not appear to preponderate to a greater extent than they do in the existing temperate flora; that the assemblage somewhat suggests American affinities; and that when we take into account deficient See also:collecting, local conditions, and the non-preservation of succulent plants, there is no reason for saying that certain other orders must have been absent. The great rarity of Monocotyledons is a common characteristic of fossil floras known only, as this one is, from leaves principally belonging to deciduous trees. With regard to suggested American affinities, it must be borne in mind that the Neocomian Angiosperms are little known except in America and in Greenland, and that we therefore cannot yet say whether families now mainly American were not formerly of world-wide distribution. We know that this was the case with some, such as Liriodendron; and in Eucalyptus we see the converse, where a genus formerly American is now confined to a far distant region. The Neocomian flora has been collected from an area extending over about 3o° of See also:latitude; but there is little evidence of any corresponding climatic change. We cannot yet say, however, that the deposits are exactly contemporaneous, and the great climatic See also:variations that have taken place in the northern hemisphere during the existence of our living flora should make us hesitate to correlate too minutely from the evidence of plants alone. The highest division of the Dakota series (known as Dakota No. I) which lies immediately beneath Upper Cretaceous strata with marine fossils, contains a flora so like that of the Tertiary deposits that only the clearest geological evidence has been considered sufficient to prove that Heer was wrong when he spoke of the plants as See also:Miocene. These highest plant-bearing strata rest, according to Lester Ward, somewhat unconformably on the Dakota No. 2; they show also a marked difference in the included plants. The genera of Dicotyledons represented are Quercus, Sassafras, Platanus, Celastrophyllum, Cissites, Viburnites. In the central parts of North America the lacustrine plant-bearing deposits are of enormous thickness, the Dakota series being followed by marine Cretaceous strata known as the See also:Colorado and See also:Montana groups, and these being succeeded conformably by a thousand feet or more of lacustrine shales, sandstones and coal-seams, belonging to the See also:Laramie series. This also contains occasional marine Upper Cretaceous fossils, as well as See also:reptiles of Cretaceous types. An extensive literature has grown up See also:relating to these Laramie strata, for owing to the Tertiary aspect of the contained plants, geologists were slow to recognize that they could be truly contemporaneous and interbedded with others yielding Cretaceous animals. In addition to this, the earlier writers included in the Laramie series many deposits now known to be of later date and truly Tertiary, and the See also:process of separation is even now only partially completed. It will be safest in these circumstances to accept as our See also:guide to the true Laramie flora the carefully compiled " Catalogue " of F. H. Knowlton. According to this catalogue, the true Laramie flora includes about 250 species, more than See also:half of which are deciduous See also:forest trees, herbaceous Dicotyledons, Monocotyledons and Cryptogams, all being but poorly represented. Among the few Monocotyledons are leaves and fruits of palms, and traces of See also:grasses and sedges. The Dicotyledons include several water-lilies, a somewhat doubtful Trapa, and many genera of forest trees still common in America. The genera best represented are Ficus (21 species), Quercus (16 species), Populus (II species), Rhamnus 9 species), Platanus (8 species), Viburnum (7 species), Magnolia (6 species), See also:Cornus (5 species), Cinnamomum (5 species), Juglans (4 species), Acer (4 species), Salix (4 species), Aralia (3 species), Rhus (3 species), Sequoia (3 species). Of trees now extinct in America, Eucalyptus and Ginkgo are perhaps the most noticeable. So large a proportion of the trees still belongs to the flora of North America that one is See also:apt to overlook the fact that among the more specialized plants some of the largest American orders, such as the See also:Compositae, are still missing from strata belonging to the Cretaceous period. The imperfection and want of continuity of the records in Europe have made it necessary in dealing with the Cretaceous floras for us to give the first place to America. But European it is now advisable to return to Europe, where cretaceous. Upper Cretaceous plants are not uncommon, and the position of the deposits in the Cretaceous series can often be fixed accurately by their close association with marine strata belonging to definite subdivisions. As these divisions of Cretaceous time will have to be referred to more than once, it will be useful to tabulate them, thus showing which plant-beds seem to be referable to each, and what are the British strata of like age. It has not yet been found possible so closely to correlate the strata of Europe with those of America, where distance has allowed geographical differences in both See also:fauna and flora to come into See also:play; therefore, beyond the references to Lower or Upper Cretaceous, no See also:classification of the American Cretaceous strata has here been given. Additional information and CommentsThere are no comments yet for this article.
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