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ENGLAND

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Originally appearing in Volume V20, Page 556 of the 1911 Encyclopedia Britannica.
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ENGLAND . See also:

FRANCE, &C. Wanting Danian Upper See also:Chalk Senonian See also:Middle Chalk Turonian See also:Lower Chalk Cenomanian Upper See also:Green-See also:sand See also:Gault See also:Albian See also:Aptian Valenginian Urgonian See also:Wealden Neocomian In the See also:continental See also:classification the deposits from the Gault downwards are grouped as Lower Cretaceous; but in See also:Great See also:Britain there is a strong break below the Gault and none above; and the Gault is therefore classed as Upper Cretaceous. The limits of the divisions in other places do not correspond, the See also:British and continental strata often being so unlike that it is almost impossible to compare them. The doubt as to the exact British See also:equivalent of the Valenginian strata of See also:Portugal, which yield the earliest Dicotyledon, has already been alluded to. The plant-bearing deposits next in See also:age, which have yielded See also:Angiosperms, appear to belong to the Cenomanian, though from See also:Westphalia a few See also:species belonging to the Cryptogams and See also:Gymnosperms, found in deposits correlated with the Gault, have been described by See also:Hosius and von der Marck. In Great Britain the whole of the Upper Cretaceous strata are of marine origin, and have yielded no See also:land-See also:plants beyond a few See also:fir-cones, See also:drift-See also:wood and rare Dicotyledonous leaves in the Lower Chalk. Most of the deposits which have yielded Angiosperms of Cretaceous age in central See also:Europe correspond in age with the See also:English Upper Chalk (Senonian), but a small Cenomanian See also:flora has been collected from the Unter Quader in See also:Moravia. Heer described from this See also:deposit at Moletein 13 genera, of which 7 are still living, containing 18 species, viz.: t See also:fern, 4 Conifers, I See also:palm, 2 See also:figs, I Credneria, 2 laurels, I Aralia, I Chondrophyllum (of uncertain See also:affinities), 2 magnolias, 2 species of Myrtaceae and a species of See also:walnut. See also:Saxony yields from strata of this See also:period at Niederschoena 42 species, de-scribed by See also:Ettingshausen. This small flora is most remarkable, for no fewer than 6 genera, containing 8 species, are referred to the Proteaceae. The Cenomanian flora of Bohemia is larger and equally See also:peculiar.

Among the See also:

Dicotyledons described by Velenovsky are the following: Credneria (5 species), Araliaceae (17 species), Proteaceae (8 species), Myrica (2 species), Ficus (5 species), Quercus (2 species), Magnoliaceae (5 species), Bombaceae (3 species), Laurineae (2 species), Ebenaceae (2 species), Verbenaceae, Conibretaceae, Sapindaceae (2 species), Camelliaceae, Ampelideae, Mimoseae, Caesalpinieae (5 species), See also:Eucalyptus (2 species), Pisonia, Phillyrea, Rhus, Prunus, Bignonia, Lower Green-sand Laurus, Salix, Benthamia. To this See also:list Bayer adds See also:Aristolochia. The Cenomanian flora of central Europe appears to be a sub-tropical one, with marked approaches to the living flora of See also:Australia. The See also:majority of its Dicotyledons belong to existing genera, but one of the most prolific and characteristic Cretaceous forms is Credneria (Fig. 3), a genus of doubtful affinities, which has been compared by different authors to the poplars; planes, limes and other orders. The Cretaceous plant-beds of Westphalia include both Upper and Lower Senonian, the two floras being very distinct. Hosius and von der Marck describe, for instance, 12 species of See also:oak from the Upper and 6 from the Lower strata, but no species is See also:common to the two. The same occurs with the figs, with 3 species above and 8 below. The 6 species of Credneria are all confined to the older deposits. In fact, not a single Dicotyledon is common to these two closely allied divisions of the Cretaceous See also:series; a circumstance not easy to explain, when we see how well the oaks and figs are represented in each. Four species of Dewalquea, a ranunculaceous genus allied to the See also:hellebore, make their See also:appearance in the Upper Senonian of Westphalia, other species occurring at See also:Aix-la-Chapelle in deposits of about the same age. The Senonian flora of the last-named See also:place, and that of Maestricht, are still only imperfectly known.

It is unnecessary to trace the See also:

variations of the Upper Cretaceous flora from point to point; but the discoveries within the See also:Arctic circle have been so surprising that See also:attention must again be called to them. Besides the Lower Cretaceous plants already mentioned, Heer has described from See also:Greenland a flora of Cenomanian age, and another belonging to the Senonian. The Cenomanian strata have yielded already 177 species, the different See also:groups being represented in these proportions: Cryptogams, 37, 30 of which are Ferns ; Cycads, 8 ; Conifers, 27 ; Monocotyledons, 8 ; See also:Ape-talons Dicotyledons, 31; other Dicotyledons, 66. The Senonian strata have yielded 118 species, 21 of which are Cryptogams, 11 Conifers, 5 Monocotyledons, 75 Dicotyledons. See also:Forest trees, especially oaks, are plentiful, and many of the species are identical with those found in Cretaceous deposits in more See also:southern latitudes. Both of these floras suggest, however, that the See also:climate of Greenland was some-what colder than that of Westphalia, though scarcely colder than warm-temperate. The Cretaceous deposits just described are followed by a series of See also:Tertiary formations, but in Europe the continuity between Cretaceous and Tertiary is not quite See also:complete. The Tertiary formations have been assigned to six periods; these are termed—Paleocene, See also:Eocene, Oligocene, See also:Miocene, See also:Pliocene, See also:Pleistocene, and each has its own botanical peculiarities. During the Paleocene period the plants were not markedly different from those of the Upper Cretaceous. Its flora is still but imperfectly known, for we are dependent on two Paleocene or three localities for the plants. There is found at Plants. Sezanne, about 6o m. See also:east of See also:Paris, an isolated deposit of calcareous tufa full of leaves, which gives a curious insight into the vegetation which flourished in Paleocene times around a See also:waterfall.

Sezanne yields Ferns in profusion, mingled with other shade-loving plants such as would grow under the trees in a moist See also:

ravine; its vegetation is comparable to that of an See also:island in the tropical seas. Monocotyledons are rare, the only ones of much See also:interest being some fragments of pandanaceous leaves. The See also:absence of Gymnosperms is noticeable. The Proteaceae are also missing; but other Dicotyledons occur in profusion, many of them being remarkable for the large See also:size of their See also:deciduous leaves. Among the flowering plants are Dewalquea, a ranunculaceous genus already mentioned as occurring in the Upper Cretaceous, and numerous living genera of forest-trees, such as occur throughout the Tertiary period, and are readily comparable with living forms. Saporta has described about seventy Dicotyledons, most of which are peculiar to this locality. The plant-bearing marls of Gelinden, near See also:Liege, contain the debris of a Paleocene forest. The trees seemed to have flourished on neighbouring chalky heights. The most abundant species of this forest were the oaks and chestnuts, of which a dozen have been collected; laurels, See also:Viburnum, See also:ivy, several Aralias, Dewalquea, a Thuja and several Ferns may be added. This flora is compared by Saporta and See also:Marion with that of southern See also:Japan. Other de-posits of this age in France have furnished plants of a more varied aspect, including myrtles, araucarias, a See also:bamboo and several See also:fan-leaved palms. Saporta points out the presence in these Paleocene deposits of certain types common, on the one See also:hand, to the See also:American Tertiary strata between the See also:Missouri and the Rocky Mountains, and on the other, to the Tertiary flora of Greenland.

The Paleocene deposits of Great Britain are of marine origin, and only yield See also:

pine-cones and fragments of Osmunda. The British Eocene and Oligocene strata yield so large a flora, and contain plant-beds belonging to so many different stages, that it is unfortunate we have still no monograph on the subject, the one commenced by Ettingshausen Eocene and and See also:Gardner in 1879 having reached no farther than ooffig(treareat t the Ferns and Gymnosperms. This deficiency Britain, makes it impossible to See also:deal adequately with the British Eocene plants, most of the material being either unpublished or needing re-examination. In the earliest Eocene plant-beds, in the See also:Woolwich and See also:Reading series, a small but interesting flora is found, which suggests a temperate climate less warm than that of earlier or of later periods. Leaves of planes are abundant, and among the plants recorded are two figs, a See also:laurel, a See also:Robinia, a Grevillea and a palm. Ferns are scarce, Ettingshausen and Gardner recording only Aneimia subcretacea and Pteris (?) Prestwichii. The only Gymnosperms determined are Libocedrus adpressa, which is See also:close to L. decurrens of the See also:Yosemite, and Taxodium europaeum. A few plants have been found in the next See also:stage, the Oldhaven beds, and among these are fig and See also:cinnamon: Gardner considers the plants to point to subtropical conditions. The See also:London See also:Clay has yielded a large number of plants, but most of the species are represented by fruits alone, not by leaves. This circumstance makes it difficult to compare the flora with that of other formations, for not only is it uncertain which leaves and fruits belong to the same plant, but there is the additional source of doubt, that different elements of the same flora may he represented at different localities. Of some plants only the deciduous leaves are likely to be preserved, whilst other succulent-leaved forms will only be known from their woody fruits. Among the 200 plants of the London Clay are no Ferns, but 6 genera of Gymnosperms—viz.

Callitris (2 species), See also:

Sequoia, Athrotaxis (?) Ginkgo, Podocar pus, Pinus; and several genera of palms, of which the tropical Nipa is the most abundant and most characteristic, among the others being fan-palms of the genera Sabal and Chamaerops. The Dicotyledons need further study. Among the fruits Ettingshausen records Quercus, See also:Liquidambar, Laurus, Nyssa, Diospyros, Symplocos, See also:Magnolia, See also:Victoria, Hightea, Sapindus, Cupania, Eugenia, Eucalyptus, Amygdalus; he suggests that the fruits of the London Clay of See also:Sheppey may belong to the same plants as the leaves found at See also:Alum See also:Bay in the Isle of See also:Wight. The next stage is represented by the Lower Bagshot See also:leaf-beds of Alum Bay. These pipeclays yield a varied flora, Ettingshausen recording 274 species, belonging to 116 genera and 63 families. Gardner, however, is unable to reconcile this estimated richness with our knowledge of the flora, and surmises that fossil plants from other localities must have been inadvertently included. He considers the flora to be the most tropical of any that has so far been studied in the See also:northern hemisphere. Its most conspicuous plants are Ficus Bowerbankii, Aralia primigenia, Comptonia acutiloba, Dryandra Bunburyi, See also:Cassia Ungeri and the fruits of Caesalpinia. The floras which it chiefly resembles are first, that of See also:Monte Bolca, and second, that of the Gres du Soissonais, which latter Gardner thinks may be of the same age, and not earlier, as is generally supposed. The See also:total number of species found at Alum Bay, according to this author, is only about 5o or 6o. To the Bagshot Sand succeeds the thick See also:mass of sands with intercalated plant-beds seen in See also:Bournemouth cliffs. Each See also:bed yields peculiar forms, the total number of species amounting to many See also:hundred, most of them differing from those occurring in the strata below.

The plants suggest a comparison of the climate and forests with those of the See also:

Malay See also:Archipelago and tropical See also:America. At one place we find drifted fruits of Nipa, at another Hightea and Anona. Other beds yield principally palms, willows, laurels, Eucalyptus or Ferns; but there are no Cycads. As showing the richness of this flora, we may mention that in the only orders which have yet been monographed, Ferns are represented by 17 species and Gymnosperms by io, though these are not the groups best represented. Gardner speaks of the Bournemouth flora as appearing to consist principally of trees or hard-wooded shrubs, comparatively few remains of the herbaceous vegetation being preserved. The higher Eocene strata of England—those above the Bournemouth Beds—are of marine origin, and yield only drifted fruits, principally fir-cones. In the volcanic districts of the See also:south-See also:west of See also:Scotland and the See also:north-east of See also:Ireland plant-beds are found intercalated between the See also:lava-flows. These also, like the lignites of Bovey Tracey, have been referred to the Miocene period, on the supposed See also:evidence of the plants; but more See also:recent discoveries by Gardner tend to throw doubt on this allocation, and suggest that, though of various ages, the first-formed of these deposits may date back to See also:early Eocene times. The flora found in See also:Mull points distinctly to temperate conditions; but it is not yet clear whether this indicates a different period from the subtropical flora of the south of England, or whether the difference depends on See also:latitude or See also:local conditions. The plants include a Fern, Onoclea hebridica, close to a living American See also:form ; four Gymnosperms belonging to the genera See also:Cryptomeria, Ginkgo, Taxus and Podocarpus; Dicotyledons of about 30 species, several of which have been figured. Among the Dicotyledons may be mentioned Platanus, Acer (?), Quercus (?), Viburnum, Alnus, Magnolia, Corylus (?), Castanea (?), Zizyphus, Populus and the See also:nettle-like Boehmeria antiqua. The absence cf the so-called cinnamon-leaves and the Smilaceae, which always enter into the See also:composition of Middle Eocene and Oligocene floras, is See also:notice-able.

The Irish strata yield two ferns; 7 Gymnosperms, Cupressus, Cryptomeria, Taxus, Podocarpus, Pinus (2 species), Tsuga; and leaves of about 25 Dicotyledons. The most abundant leaf, according to Gardner, does not seem distinct from Celastrophyllum Benedeni, of the Paleocene strata of Gelinden; a See also:

water-See also:lily, Nelumbium Buchii, occurs also in Oligocene beds on the See also:Continent; the species of MacClintockia (fig. 4) is found both in the Arctic floras and at Gelinden. Among the other plants are an See also:alder, an oak and a doubtful cinnamon. Leaving these Scottish and Irish deposits of doubtful age, we find in the See also:Hampshire See also:Basin a thick series of fluviatile, lacustrine and marine deposits undoubtedly of Lower and Middle Oligocene date. Their flora is still a singularly poor one, though plants have been obtained at many different levels; they perhaps indicate a somewhat cooler climate than that of the Bournemouth series. Among the more abundant plants are nucules of several species of Chara, and drifted fruits and seeds of water-lilies, of Folliculites (now generally referred to Stratiotes) and of Limnocarpus (allied to Potamogeton) ; there is little else mixed with these. Other seams are full of the twigs and cones of Athrotaxis, a Conifer now confined to See also:Tasmania. Ferns are represented by Gleichenia, Lygodium and Chrysodium Lanzaeanum, which last has a very wide range in See also:time; Monocotyledons, by a Sabal and a See also:feather-palm, as well as by the two aquatic genera above mentioned; Gymnosperms. by the See also:extinct araucarian genus Doliostrobus, by rare pine-cones, and by Athrotaxis. Dicotyledonous leaves are not plentiful, the genera recorded being See also:Andromeda, Cinnamomum, Zizyphus, Rhus, Viburnum. The See also:lignite deposits and See also:pipe-See also:clays of Bovey Tracey in See also:Devon, referred by Heer and See also:Pengelly to the Miocene period, were considered by Gardner to be of the same age as the Bournemouth beds (Middle Eocene). Recent researches show, however, that Heer's view was more nearly correct.

The flora of Bovey is like that of the lignite of the Wetterau, which is either highest Oligocene or lowest Miocene. Several species of Nyssa are common to the two districts, as are a climbing palm, two vines, a magnolia, &c. The common See also:

tree at Bovey is Sequoia Couttsiae, which probably See also:grew in profusion in the sheltered valleys of See also:Dartmoor, close to the See also:lake. Above these strata in Great Britain there is a complete break, no species of plant ranging upwards into the next fossiliferous See also:division. central and that met with in other See also:geographical regions and in Southern France. other latitudes. For this study it will be most convenient to take next south and central France, for in that See also:area can be found a series of plant-bearing strata in which is preserved a nearly continuous See also:history of the vegetation from Upper Eocene down to Pliocene. The See also:account is taken mainly from the writings of Saporta. The See also:gypsum-deposit of Upper Eocene date at Aix in See also:Provence commences this series, and is remarkable for the variety and perfect preservation of its organic remains. Among its Gymnosperms are numerous Cupressineae of See also:African See also:affinity belonging to the genera Callitris and Widdringtonia, and a See also:juniper close to one indigenous in See also:Greece. Fan-palms, several species of See also:dragon-tree and a See also:banana, like one living in See also:Abyssinia, represent the more peculiar Mono-cotyledons. Among the noticeable Dicotyledons are the Myricaceae, Proteaceae, Laurineae, Bombax, the Judas-tree, See also:Acacia, See also:Ailanthus, while the most plentiful forms are the Araliaceae. Willows and poplars, with a few other plants of more temperate regions, are found rarely at Aix, and seemingly point to casual introduction from surrounding mountains.

In a See also:

general way, spiny plants, with stiff branches and dry and coriaceous leaves, dominate theflora, as they now do in Central See also:Africa, to which region on the whole Saporta considers the flora to be most allied. The succeeding Oligocene flora appears to. be more characterized by a See also:gradual replacement of the Eocene species by allied forms, than by any marked See also:change in the assemblage or in the See also:climatic conditions. It forms a perfectly gradual transition to the still newer Miocene period, the newer species slowly appearing and increasing in number. Saporta considers that in central and southern Europe the alternate dry and moist See also:heat of the Eocene period gave place to a climate more equally and more universally humid, and that these conditions continued without material change into the succeeding Miocene stage. Among the types of vegetation which make their appearance in Europe during the Oligocene period may be mentioned the Conifers Libocedrus salicornioides, several species of Chamaecyparis and Sequoia, Taxodium distichum and Glyptostrobus europaeus. The palms include Sabal haeringiana, S. See also:major and Flabellaria. Among the Myricaceae several species of Comptonia are common. These new-corners are all of American type. Aquatic plants, especially water-lilies, are abundant and varied; the See also:soil-dry Callitris and Widdringtonia become scarce. Though we do not propose to deal with the other See also:European localities for Eocene and Oligocene plants, there is one See also:district to which attention should be See also:drawn, on account of the exceptional See also:state of preservation of the specimens. plantser m Amb On the Baltic shores of See also:Prussia there is found a quantity of See also:amber, containing remains of See also:insects and plants. This is derived from strata of Oligocene age, and is particularly valuable because it preserves perfectly various soft parts of the plants, which are usually lost in fossil specimens. The tissues, in fact, are preserved just as they would be in .See also:Canada See also:balsam.

The amber yields such things as fallen See also:

flowers, perfect catkins of oak, See also:pollen grains and See also:fungi. It enables us to determine accurately orders and genera which otherwise are unknown in the fossil state, and it thus See also:aids us in forming a truer See also:idea of the flora of the period than can be formed at any locality where the harder parts alone are recognizable. No doubt this amber flora is still imperfectly known, but it is valuable as giving a See also:good idea of the vegetation, during Oligocene times, of a mixed wood of pine and oak, in which there is a mixture of herbaceous and woody plants, such as would now be found under similar conditions. The plants of which the floral See also:organs or perfect fruits are pre-served include the amber-bearing Pinus succinifera, Smilax, See also:Phoenix, the spike of an aroid, II species of oak, 2 of See also:chestnut, a See also:beech, Urticaceae, 2 cinnamons and Trianthera among the Lauraceae, representatives of the Cistaceae, Ternstroemiaceae, Dilleniaceae (3 species of Hibbertia), See also:Geraniaceae (See also:Geranium and Erodium), Oxalidaceae, Acer, Celastraceae, Olacaceae, Pittosporaceae, Ilex (2 species), See also:Euphorbiaceae, See also:Umbelliferae (Chaerophyllum), See also:Saxifragaceae (3 genera), Hamamelidaceae, See also:Rosaceae, Connaraceae, See also:Ericaceae (Andromeda and Clethra), Myrsinaceae (3 species), See also:Rubiaceae, Sambucus (2 species), Santalaceae, Loranthaceae (3 species). We here discover for the first time various living families and genera, but there is still a noticeable absence of many of our most prolific existing groups. Whether this deficiency is accidental or real time will show. The Miocene flora, which succeeds to that just described, is well represented in Europe; but till recently there has been an unfortunate tendency to refer Tertiary floras of all Miocene. See also:dates to the Miocene period, unless the See also:geological position of the strata was so clear as obviously to forbid this See also:assignment. Thus plant-beds in the See also:basalt of Scotland and Ireland were called Miocene; and in the Arctic regions and in North America even plant-beds of Upper Cretaceous age were referred to the same period. The See also:reason for this was that some of the first Tertiary floras to be examined were certainly Miocene, and, when these plants had been studied, it was considered that somewhat similar assemblages found elsewhere in deposits of doubtful geological age must also be Miocene. For a See also:long time it was not recognized that changes in the marine See also:fauna, on which our geological classification mainly depends, correspond scarcely at all with changes in the land plants. It was not suspected, or the fact was ignored, that the break between Cretaceous and Tertiary—made so conspicuous by striking changes in the aquatic animals—had little or no importance in botanical history. It was not realized that an Upper Cretaceous flora needed See also:critical examination to distinguish it from one of Miocene age, and that the two periods were not characterized by a Space will not allow us to deal with the numerous scattered deposits which have yielded Tertiary plants.

It will be more to the purpose to take distant areas, where the See also:

order of the strata is clear, and compare the See also:succession of the floras with sweeping change of generic type, such as took place among the marine invertebrates. It may appear absurd to a geologist that any one could See also:mistake a Cretaceous flora for one of Miocene date, since the marine animals are completely different and the See also:differences are striking. In the See also:case of the plants, however, the Tertiary generic types in large See also:part appeared in Upper Cretaceous times. Few or no extinct types are to be found in these older strata—there is nothing among the plants equivalent to the unmistakably extinct See also:Ammonites, Belemnites, and a hundred other groups, and we only meet with See also:constant variations in the same genus or See also:family, these variations having seldom any obvious relation to phylogeny. The Miocene period is unrepresented by any deposits in Great Britain, unless the Bovey lignite should belong to its earliest stage; we will therefore commence with the best known region—that of central Europe and especially of See also:Switzerland, whence a prolific flora has been collected and described by See also:Oswald Heer. The Miocene lacustrine deposits are contained in a number of silted-up lake-basins, which were successively formed and obliterated during the uprise of the See also:Alps and the continuous folding and bending of the See also:earth's crust which was so striking a feature of the period. These undulations tended to transform valleys into chains of lakes, into which the plants and animals of the surrounding area See also:fell or were washed. We thus find preserved in the Upper Miocene lacustrine deposits of Switzerland a larger flora than is known from any other period of similar length; in fact, an See also:analysis of its composition suggests that the Miocene flora of Switzerland must have been both larger and more varied than that now living in the same See also:country. The best known locality for the Upper Miocene plants is Oeningen, on the Lake of See also:Constance, where have been collected nearly 50o species of plants, the total number of Miocene plants found in Switzerland being stated to be now over 900. Among the characteristics of this Miocene flora are the large number of families represented, the marked increase in the deciduous-leaved plants, the gradual decrease in the number of palms and of tropical plants, and the replacement of these latter by Mediterranean or North American forms. According to Heer, the tropical forms in the Swiss Miocene agree rather with See also:Asiatic types, while the subtropical and temperate plants are allied to forms now living in the temperate See also:zone in North America. Of the 920 species described by Heer, 114 are Cryptogams and 806 flowering plants.

Mosses are extremely rare, Heer only describing 3 species. Vascular Cryptogams still include one or two large horsetails with stems over an See also:

inch thick, and also 37 species of Fern, amongst the most interesting of which are 5 species belonging to the climbing Lygodium, a genus now living in See also:Java. The number of Ferns is just equal to that now found in Switzerland. Cycads are only represented by fragments of two species, and this seems to be the last appearance of Cycads in Europe. The Coniferae include no fewer than 94 species of Cupressineae and 17 of Abietineae, including several species of Sequoia. Monocotyledons form one-See also:sixth of the known Miocene flora, 25 of them being See also:grasses and 39 sedges; but most of these need further study, and are very insufficiently characterized. Heer records one species of See also:rice and four of See also:millet. Most of the other Monocotyledons See also:call for little remark, though among them is an See also:Iris, a Bromelia and a See also:ginger. Smilax, as in earlier times, was common. Palms, referred to i i species, are found, though they seem to have decreased in abundance; of them 7 are fan-palms, the others including Phoenicites—a form allied to the date—and a trailing palm, Calamopsis, allied to the canes and rattans. Among the Dicotyledons, the See also:Leguminosae take the first place with 131 species, including Acacia, Caesalpinia and Cassia, each represented by several forms. The occurrence of 90 species of Amentaceae shows that, as the climate became less tropical, the relative proportion of this See also:group to the total flora increased.

See also:

Evergreen oaks are a marked characteristic of the period, more than See also:half the Swiss species being allied to living American forms. Fig-trees referred to 17 species occur, all with undivided leathery leaves; one is close to the See also:banyan, another to the indiarubber-tree The Laurineae were plentiful, and include various true laurels, camphor-trees, cinnamon, Persea and Sassafras. The Proteaceae, according to Heer, are still common, the Australian genera Hakea, Dryandra, Grevillea and See also:Banksia, being represented. Amongst gamopetalous plants several of our largest living families, including Campanulaceae, See also:Labiatae, See also:Solanaceae and See also:Primulaceae, are still missing; and of Boragineae, Scrophularineae, Gentianeae and See also:Caprifoliaceae there are only faint and doubtful indications. The Cotnpositae are represented by isolated fruits of various species. See also:Twining lianas are met with in a species of Bignonia; Umbelliferae See also:Ranunculaceae and See also:Cruciferae, are represented by a few fruits. These families, however, do not appear to have had anything like their See also:present importance in the temperate flora, though, as they are mainly herbaceous plants with fruits of moderate hardness, they may have decayed and See also:left no trace. The American Liriodendron still flourished in Europe. Water-lilies of the genera Nymphaea and Nelumbium occur. Maples were still plentiful, 20 specieshaving been described. Rosaceae are rare, Crataegus, Prunus and Amygdalus, being the only genera recorded. It is obvious that many of these Swiss Miocene plants will need more close study before their specific characters, or even their generic position, can be accepted as thoroughly made out; still, this will not affect the general composition of the flora, with its large proportion of deciduous trees and evergreens, and its noticeable deficiency in many of our largest living families.

From Europe it will be convenient to pass to a distant region of similar latitude, so that we may see to what extent botanical provinces existed in Eocene and Oligocene times. It Tertiary so happens that the interior of temperate North of North America is almost the only region outside Europe in Amer" which a series of plant-bearing strata give a connected history of these periods, and in which the plants have been collected and studied. It is unfortunately still very difficult to correlate even approximately the strata on the two sides of the See also:

Atlantic, and there is great doubt as to what strata belong to each division of the Tertiary period even in different parts of North America. This difficulty will disappear as the strata become better known; but at present each of the silted-up lakes has to be studied separately, for we cannot expect so close a See also:correspondence. in their faunas and floras as is found in the more crowded and smaller basins in central Europe. Perhaps the most striking characteristic of the Tertiary floras of North America, as distinguished from those of Europe, is the greater continuity in their history and greater connexion with the existing flora of the same regions. This difference is readily explained when we remember that in Europe the See also:main barriers which stop See also:migration, such as the Alps and the Mediterranean, run east and west, while in America the only barriers of any importance run north and south. In consequence of this peculiarity, climatic or orographic changes in Europe tend to drive animals and plants into a cul de See also:sac, from which there is no See also:escape; but in America similar climatic waves merely cause the species alternately to See also:retreat and advance. .This difficulty in migration is probably the reason why the existing European flora is so poor in large-fruited trees compared with what it was in Miocene times or with the existing flora of North America. In America the contrast between the Eocene forests and those now living is much less striking, and this fact has led to the wrong See also:assumption that the present American flora had its origin in the American continent. Such a conclusion is by no means warranted by the facts, for in Tertiary times, as we have seen, the European flora had a distinctly " American " facies. Therefore the so-called American forms may have originated in the Old See also:World, or more probably, as Saporta suggests, in the polar regions, whence they were driven by the increase of See also:cold southwards into Europe and into America. The American Tertiary flora is so large, and the See also:geology of the deposits is so intricate, that it is out of the question to discuss them more fully within the limits of this See also:article.

We may point out, however, that the early Tertiary floras seem to indicate a much closer connexion and a greater community of species than is found between the existing plants of Europe and America. Or, rather, we should perhaps say that See also:

ancient floras suggest recent dispersal from the place of origin, and less time in which to vary and become modified by the loss of different groups in the two continents. Geographical provinces are certainly indicated by the Eocene flora of Europe and America, but these are less marked than those now existing. If we turn to a more isolated region, like Australia, we find a Lower Eocene flora distinctly related to the existing flora of Australia and not to that of other continents. See also:Australasia had then as now a peculiar flora of its Australia. own, though the former wide dispersal of the Proteaceae and Myrtaceae, and also the large number of Amentaceae then found in Australia, make the Eocene plants of Europe and Australia much less unlike than are the present floras. Within the Arctic circle a large number of Tertiary plants have been collected. These were described by Heer, who referred them to the Miocene period; he recognized, Arctic in fact, two periods during which Angiosperms Regions. flourished within the Arctic regions, the one Upper Cretaceous, the other Miocene. To this view of the Miocene age of the plant-bearing strata in Greenland and See also:Spitsbergen there are serious objections, which we will again refer to when the flora has been described. The Tertiary flora of Greenland is of great interest, from the extremely high latitude at which the plants flourished, See also:thirty of the species having been collected so far north as See also:lat. 81°. Taking first this most northerly locality, in See also:Grinnell Land, we find the flora to comprise 2 horsetails, 11 Conifers (including the living Pinus Abies), 2 grasses, a sedge, 2 poplars, a See also:willow, 2 birches, 2 hazels, an See also:elm, a Viburnum, a water-lily, and a See also:lime.

Such an assemblage at the present See also:

day would suggest a latitude quite 25° farther south; but it shows decidedly colder conditions than any of the European Eocene, Oligocene, or Miocene strata. From lat. 78° in Spitsbergen Heer records 136 species of fossil plants. More to the south, at Disco Island in lat. 70°, the Tertiary wood seem to have been principally composed of planes and Sequoias; but a large number of other genera occur, the total number of plants already recorded being 137. From various parts of Greenland they now amount to at least 280. Among the plants from Disco, more than a See also:quarter are also found in the Miocene of central Europe. The plants of Disco include, besides the See also:plane and Sequoia, such warm-temperate trees as Ginkgo, oak, beech, See also:poplar, See also:maple, walnut, lime and magnolia. If these different deposits are contemporaneous, as is not improbable, there is a distinct change in the flora as we move farther from the See also:pole, which suggests that difference of latitude then as now was accompanied by a difference in the flora. But if this See also:process is continuous from latitude to latitude, then we ought not to look for a flora of equivalent age in the warm-temperate Miocene deposits of central Europe, but should rather expect to find that the temperate plants of Greenland were contemporaneous with a tropical flora in central Europe. As Mr Starkie Gardner has pointed out, it does not seem reasonable to assume that the same flora could have ranged then through 40° of latitude; it is more probable that an Eocene temperate flora found in the Arctic regions travelled south-wards as the climate became cooler, till it became the Miocene temperate flora of central Europe. Mr Gardner suggests, therefore, that the plant-beds of Greenland and Spitsbergen represent the period of greatest heat, and are therefore wrongly referred to the Miocene.

At present the evidence is scarcely sufficient to decide the question, for if this view is right, we ought to find within the Arctic circle truly Arctic floras equivalent to the cool Lower Eocene and Miocene periods; but these have not yet been met with. A steady decrease of temperature marked the Pliocene period throughout Europe, and gradually brought the climatic See also:

con- Piiocene. ditions into correspondence with those now existing, till towards the end of the period neither climate nor See also:physical See also:geography differed greatly from those now existing. Concurrently with this change, the tropical and extinct forms disappeared, and the flora approached more and more nearly to that now existing in the districts where the fossil plants are found, though in the older deposits, at any See also:rate, the geographical See also:distribution still differed considerably from that now met with. At last, in the latest Pliocene strata (often called " pre-Glacial ") we find a flora consisting almost entirely of existing species belonging to the Palaearctic regions, and nearly all still living in the country where the fossils are found. This flora, however, is associated with a fauna of large mammals, the majority of which are extinct. The plants of the Older Pliocene period are unknown in Great Britain, and little known throughout Europe except in central France and the Mediterranean region. The forests of central France during this See also:epoch showed, according to Saporta, a singular admixture of living European species, with trees now characteristic of the See also:Canary Isles and of North America. For instance, of the living species found at Meximieux, near See also:Lyons, one is American, eight at least belong to the Canaries (six being characteristic of those islands), two are Asiatic, and ten still live in Europe. Taking into account, however, the closest living See also:allies of the fossil plants, we find about equal affinities with the floras of Europe, America, and See also:Asia. There is also a decided resemblance to the earlier Miocene flora. Among the more interesting plants of this deposit may be mentioned Torreya nucifera, now See also:Japanese; an evergreen oak close to the common Quercus Ilex; Laurus canariensis, Apollonias canariensis, Persea carolinensis, and Ilex canariensis; See also:Daphne pontica (a plant of Asia See also:Minor); a species of See also:box, scarcely differing from the English, and a bamboo. To this epoch, or perhaps to a stage slightly later, and not to the Newer Pliocene period, as is generally supposed, should probably be referred the lignite deposits of the Val d'See also:Arno.

This lignite and the accompanying leaf-bearing clays underlie and are apparently older than the strata with Newer Pliocene mammals and See also:

mollusca. The only mammal actually associated with the plants appears to be a species of See also:tapir, a genus which in Europe seems to be characteristically Miocene and Older Pliocene. The plants of the Val d'Arno have been described by See also:Ristori; they consist mainly of deciduous trees, a large proportion of which are known Miocene and early Pliocene forms, nearly all of them being extinct. A markedly upland See also:character is given to the flora of this valley through the abundance of pines (9 species) and oaks (16 species) which it contains; but this peculiarity is readily accounted for by the steep slopes of the See also:Apennines, which everywhere surround and dominate the old lake-basin. Among the other noticeableplants may be mentioned Betula (3 species), Alnus (2 species), Carpinus, Fagus (4 species), Salix (4 species), Populus (2 species), Platanus, Liquidambar, Planera, Ulmus (2 species), Ficus (2 species), Persoonia, Laurus (5 species), Persea, Sassafras, Cinnamomum (5 species), Oreodaphne, Diospyros (2 species), Andromeda, Magnolia, Ater (3 species), Sapindus, Celastrus (2 species), Ilex (4 species), Rhamnus (3 species), Juglans (5 species), Carya (2 species), Rhus, Myrtus, Crataegus, Prunus, Cassia (3 species). These plants suggest a colder climate than that indicated by the plants of Meximieuxthey might, therefore, be thought to belong to a later period. The difference, however, is probably fully accounted for when we take into See also:consideration the biting winds still See also:felt in See also:spring in the valley of the Arno, and the probable large admixture of plants washed down from the mountains above. Somewhat later Pliocene deposits in the Val d'Arno, as well as the tuffs associated with the Pliocene volcanoes in central France, yield plants of a more See also:familiar type, a considerable proportion of them still living in the Mediterranean region, though some are only now found at distant localities, and others are extinct. The flora, however, is essentially Palaearctic, American and Australian types having disappeared. A somewhat later Pliocene flora is represented by the plants found at Tegelen, near Venloo, on the See also:borders of the See also:Netherlands and See also:Germany. This deposit is of especial interest for the See also:light it throws on the origin of the existing flora of Britain. The Tegelen plants are mainly north European; but there occur others of central and south Europe, and various See also:exotic and extinct forms, nearly all of which, however, belong to the Palaearctic region, though some may now be confined to widely separated parts of it.

For instance, Pterocarya caucasica does not grow nearer than the See also:

Caucasus, where it is associated with the See also:wild vine—also found at Tegelen; Magnolia Kobus is confined to the north island of Japan; another species of Magnolia cannot be identified and may be extinct. An extinct water-lily, Euryale limburgensis, belongs to a See also:monotypic genus now confined to See also:Assam and See also:China; an extinct sedge, Dulichium vespiforme, belongs to a genus only living in America, though the only living species once flourished also in See also:Denmark; an extinct species of water-See also:aloe (Stratiotes elegans) makes a third genus, represented only by a single living species, which was evidently better represented in Pliocene times. A large proportion of the plants, however, may still be found living in See also:Holland and Britain; but there is a singular scarcity of Composites, though this order is fairly well represented in British strata of slightly later date. The latest Pliocene, or pre-Glacial, flora of northern Europe is best known from the See also:Cromer Forest-bed of See also:Norfolk and See also:Suffolk, a fluvio-marine deposit which lies beneath the whole of the Glacial deposits of these counties, and passes downwards into the See also:Crag, many of the animals actually associated with the plants being characteristic Pliocene species which seem immediately afterwards to have been exterminated by the increasing cold. The plants contained in the Cromer Forest-bed, of which about 15o species have now been determined, fall mainly into two groups—the forest-trees, and See also:marsh and aquatic plants. We know little or nothing at present of the upland plants, or of those of dry or chalky soils. Forest trees are well represented; they are, in fact, better known than in any of the later English deposits. We find the living British species of Rhamnus, maple, sloe, See also:hawthorn, See also:apple, See also:white-See also:beam, guelder-See also:rose, cornel, elm, See also:birch, alder, See also:hornbeam, See also:hazel, oak, beech, willow, See also:yew and pine, and also the spruce. This is an assemblage that could not well be found under conditions differing greatly from those now holding in Norfolk; there is an absence of both Arctic and south European plants. The variety of trees shows that the climate was mild and moist. Among the herbaceous plants we find, mingled with a number that still live in Norfolk, Hypecoum procumbens, the water-chestnut (Trapa natans), and Najas minor, none of which is now British. On the Norfolk See also:coast another thin plant-bed occurs locally above the Forest-bed and immediately beneath the See also:Boulder Clay.

This deposit shows no trace of forest-trees, but it is full of remains of Arctic mosses, and of the See also:

dwarf willow and birch ; in See also:short, it yields the flora now found within the Arctic circle. The incoming of the Glacial epoch does not appear to have been accompanied by any See also:acclimatization of the Dlants—the species belonging to temperate Europe were locallymeistocene. exterminated, and Arctic forms took their places. The same Arctic flora reappears in deposits immediately above the highest Boulder Clay, deposits formed after the See also:ice had passed away. These fossil Arctic plants have now been found as far south as Bovey Tracey in See also:Devonshire, where Pengelly and Heer discovered the See also:bear-See also:berry and dwarf birch; London, where also Betula nana occurs; and at Deuben in Saxony, which lies nearly as far south as lat. 5o°, but has yielded to See also:Professor Nathorst's researches several Arctic species of willow and See also:saxifrage. The cold period, however, was not continuous, for both in Great Britain and on the continent of Europe, as well as in Canada, it was broken by the recurrence of a milder climate and the reappearance of a flora almost identical with that now living in the same regions. This " inter-Glacial " flora, though so like that now found in the district, has interesting peculiarities. In England, for instance, it includes Acer monspessulanum, a southern maple which does not now extend nearer than central Europe, and See also:Cotoneaster Pyracantha; also Najas graminea and N. minor, both southern forms not now native of Britain. Brassenia peltata, a water-lily found in the warmer regions almost throughout the world, except in Europe, occurs abundantly in north Germany, but not in Great Britain. Similar inter-Glacial deposits in See also:Tirol contain leaves of See also:Rhododendron ponticuin. Space will not permit us to enter into any full discussion of the recurrence of Glacial and inter-Glacial periods and the See also:influence they may have had on the flora. It is evident, how-ever, that if climatic alternations, such as those just described, are part of the normal routine that has gone on through all geological periods, and are not merely confined to the latest, then such changes must evidently have had great influence on the See also:evolution and geographical distribution both of species and of floras.

Whether this was so is a question still to be decided, for in dealing with extinct floras it is difficult to decide, except in the most general way, to what climatic conditions they point. We seem to find indications of long-period climatic oscillations in Tertiary times, but none of the sudden invasion of an Arctic flora, like that which occurred during more recent times. It should not be forgotten, however, that an Arctic flora is mainly distinguishable from a temperate one by its poverty and dwarfed vegetation, its deciduous leaves and small fruits, rather than by the occurrence of any characteristic genera or families. Careful and long-continued study would therefore be needed before we could say of any extinct dwarfed flora that it included only plants which could withstand Arctic conditions. AuruoRlrlES.—H. Conwentz, Monographie der baltiSchen Bernsteinbaume (See also:

Danzig, 1890), See also:Die Flora See also:des Bernsteins, vol. ii. (1886) ; See also:Sir W. See also:Dawson, Papers on the Cretaceous Plants of British North America, Trans. See also:Roy. See also:Soc. Canada (1883-1896); C. von Ettingshausen, Die Kreideflora von Niederschona in Sachsen," Sift. le. Akad.

Wiss. Wien, math.-nat. Cl., vol. lv., Abth. i. (1867) ; " See also:

Report on . . . Fossil Flora of Sheppy," Proc. Roy. Soc. See also:xxix. 388 (1879) ; Report on . Fossil Flora of Alum Bay," ibid. See also:xxx. 228 (188o) ; C. von Ettingshausen and J. S.

Gardner, " Eocene Flora," vols. i. and ii., Palaeont. Soc. (1879-1886) ; W. M. See also:

Fontaine, " The See also:Potomac or Younger Mesozoic Flora," U.S. Geological Survey, Monograph xv. (1889) ; J. S. Gardner, Flora of Alum Bay, in " Geology of the Isle of Wight," Mem. Geol. Survey (2nd ed., 1889); H. R.

Goeppert and A. Menge, Die Flora des Bernsteins and ihre Beziehungen zur Flora der Tertiarformation and der Gegenwart, vol. i. (Danzig, 1883); 0. Heer, Flora tertiaria ilehetiae (3 vols., See also:

Winterthur, 1855-1859) ; Flora fossilis arctica (7 vols., See also:Zurich, 1868-1883), " Beitrage zur Kreideflora,—(1) Flora von Moletein in Mahren," New Denkschr. allgem. schweiz. Gesell. Naturwiss., vol. See also:xxiii. mem. 22 (Zurich, '869-1872); Primaeval World in Switzerland (2 vols., '876); F. H. Knowlton, ' See also:Catalogue of the Cretaceous and Tertiary Plants of North America," See also:Bull. U.S. Geol. Survey (No.

152, 1898), " Flora of the See also:

Montana Formation," ibid., No. 163 (19oo); Krasser, " Die fossile Kreideflora von Kunstadt in Mahren,' See also:Beit. paleont. Geol. Oesterreich-Ungarns, Bd. v. Hft. 3 (1896); See also:Leo. Lesquereux, " Contributions to the Fossil Flora of the Western Territories," See also:Rep. U.S. Geol. Survey of the Territories, vols. vi., vii., viii. (1877-1883), " The Flora of the Dakota Group," U.S. Geological Survey, Monograph xvii.

(1891); Meschinelli and Sduinabol, Flora tertiaria italica (1892); this See also:

book contains a full bibliography See also:relating to the Fossil Flora of See also:Italy; J. S. See also:Newberry, "The Flora of Amboy Clays," U.S. Geological Survey, Monograph See also:xxvi. (1895); Hosius and von der Marck, " Die Flora der westphalischen Kreideformation," Palaeontographica, vol. xxvi. (188o), and supplement in ibid. vol. xxxi. (1883) ; A. G. Nathorst, " Glacialflora in Sachsen, am aussersten Rande des nordischen Diluviums," Kongl. Vetenskaps-Akad. Forh., p. 519 (1894) ; See also:Clement See also:Reid, " Pliocene Deposits of Britain," Mem.

Geol. Survey (189o), Origin of the British Flora (1899); C. and E. M. Reid, " The Fossil Flora of Tegelen-sur-See also:

Meuse, near Venloo, in the See also:Province of See also:Limburg," Verh. Kon. Akad. Wetensch. See also:Amsterdam, 2e See also:Sect. DI. xiii. No. 6 (1907) ; " On the Pre-Glacial Flora of Britain," Journ. Linn.

Soc. (See also:

Botany), xxxviii. 206-227 (1908); G. de Saporta, Prodrome dune flore fossile des Travertins anciens de Sezanne," Mem. soc. geol. France, 2nd series, vol. viii. p. 28? (1868); " Recherches sur See also:les vegetaux fossiles de Meximieux, Archiv. See also:Mus. hist. nat. See also:Lyon, i. 131 (1876); Monde des plantes avant l'apparition de l'homme (1879) ; ' Etudes sur la vegetation du sud-est de la France a 1'epoque tertiare," See also:Ann. sci. nat. (1862-1888); Flore fossile du Portugal (See also:Lisbon, 1894) ; G. de Saporta and A. F. Marion, " Essai sur 1'etat de la vegetation a 1'epoque des marnes heersiennes de Gelinden," Mem. tour. acad. See also:ray. belgique, vol. See also:xxxvii.

No. 6 (1873), and vol. xli. No. 3 (1878); J. Velenovsky, " Die Flora der bohmischen Kreideformation," in Beitrage zur Paleontologie Oesterreich-Ungarns and des Orients, vols. ii.-v. (1881-1885); Lester F. See also:

Ward, " Synopsis of the Flora of the See also:Laramie Group," 6th Report U.S. Geological Survey, pp. 399-558 (1885) ; " The Geographical Distribution of Fossil Plants," 8th Report U.S. Geological Survey, pp. 663-96o (1889) ; " The Potomac Formation," 75th Report U.S. Geological Survey, pp.

307-398 (1895) ; " Some Analogies in the Lower Cretaceous of Europe and America," r6th Report U.S. Geological Survey, Pt. I., pp. 462-542 (1896) ; " The Cretaceous Formation of the See also:

Black Hills as indicated by the Fossil Plants," 79th Report U.S. Geological Survey, Pt. II., pp. 521-946 (1899). (C.

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