in Ae IAN SAAATO NN iH ) 4 \ \ a Ry NN bs) LAN AUN Hh tis Wit thet Peer the hs apetsde Aah NSA TEMS CNT BAY mR i irda Pat Rear 54 dil vires ep ig Las Ua ede 4 Pidictyontr ath oe ats oe Tra i rad 4 ot mets Be is a ia ih 9 ie ALBERT R. MANN LIBRARY AT CORNELL UNIVERSITY 3 1924 057 348 165 Cornell University Library The original of this book is in the Cornell University Library. There are no known copyright restrictions in the United States on the use of the text. http://www.archive.org/details/cu31924057348165 THE NATURAL HISTORY OF PLANTS THEIR FORMS, GROWTH, REPRODUCTION, AND DISTRIBUTION FROM THE GERMAN OF ANTON KERNER von MARILAUN PROFESSOR OF BOTANY IN THE UNIVERSITY OF VIENNA BY F. W. OLIVER, M.A,, D.Sc. QUAIN PROFESSOR OF BOTANY IN UNIVERSITY COLLEGE, LONDON WITH THE ASSISTANCE OF MARIAN BUSK, BSc. anp MARY F. EWART, BSc. WITH ABOUT 1000 ORIGINAL WOODCUT ILLUSTRATIONS AND SIXTEEN PLATES IN COLOURS HALF-VOLUME IV. NEW YORK HENRY HOLT AND COMPANY 1895 A. g05 se CONTENTS OF HALF-VOLUME IV. LIST OF ILLUSTRATIONS. PuateE XV. FRONDOSE AND FRutIcOosE LICHENS, - - - » XVI. Evcatyprus GRovE AND GRASS-TREES IN AUSTRALIA, Illustrations in the Text—Fig. 357 to Fig. 482. THE HISTORY OF SPECIES (Continued), 2, Alteration in the Form of Species (Continued)— The Influence of Mutilation on the Form of Plants, - - - Alteration of Form by Parasitic Fungi, - Alteration of Form by Gall-producing Insects,- - - - The Genesis of New Forms as a result of Crossing, 3. The Origin of Species— The Genesis of New Species, - BR So us Derivation of Existing Species, ‘- - : 2 The Subdivisions of the Vegetable Kingdom, - 5 4, The Distribution of Species— The Distribution of Species by Offshoots, - The Dispersion of Species by Means of Fruits and Seeds,- —- Limits of Distribution, 3 = = Plant Communities and Floras, - < s 5. The Extinction of Species, - - e.g to face PAGE 694 730 514 518 527 554 576 595 600 790 833 878 885 899 ERRATA. Vou. I. Plate I., fly-sheet, q, for Cress read Summer Savory (Satureja hortensis). p. 336, line 15, for Daffodil read Lily. p. 547, line 16, for 16°1° read — 16°1°. p. 663, line 27, for repens read reptans. p. 685, line 24, delete cownter-. » line 28, for clockwise read cownter-clockwise. Vou. II. Pp. 69, line 4, for Lycopodiacece read Lycopodiales. p. 89, line 12, for 213’ and 213 ? read 213? and 213 *, p. 120, line 7 from bottom, for acer read acris. p. 121, line 1, for Leguminous read Oruciferous. p. 280, line 2 from bottom, for Lentibulacee read Lentibularincee. p. 612, line 11 from the bottom, for calcination read calcification. p. 633, line 1, after though insert not. p. 641, line 12, for not read -net. p- 648, line 7, for the read some. » line 16, for allimetes read akinetes. p. 696, line 25 from bottom, for Antherocerotacee read Anthocerotacew. p. 698, line 1, for Antherocerot read Anth ti p. 704, line 14, for Pteridophyte read Pteridophyta. INFLUENCE OF THE SUBSTRATUM. 497 Under these circumstances it is a matter of indifference whether 10 per cent or only traces of lime or silica can be demonstrated in the soil, and the hypothesis that plant-species which grow on limestone fail to grow on slate because they are not able to supply their need of calcium, or that the plants growing on slate cannot flourish on limestone mountains because they cannot obtain the necessary amount of silica, must be abandoned, as well as the assumption that these substances. when absorbed as food serve as a stimulus to change of form. I strongly supported this latter hypothesis at the time, and thought I should be able to strengthen and confirm it by careful cultural experiments. Seeds of several species which demand lime were sown in soil containing hardly perceptible quanti- ties of lime, and the seedlings were watered with water devoid of calcium; in another place seeds of species demanding a silica-containing substratum were placed in soil which contained much limestone, and the seedlings were watered with lime- water. At first it seemed as if an alteration of form had actually taken place in -some individuals. But this was a mistake, or rather, the alteration only consisted in the greater or less luxuriance of the foliage, lengthening or shortening of the stem, abundant or scanty development of flowers and the like. But no actual change of form which would be retained by their descendants could be obtained. The species of plants accustomed to lime, grown on a soil devoid of lime, presented a miserable appearance, with scanty flowers which ripened only a few seeds, whilst the silica-demanding species grown on lime-containing soil soon withered and died without flowering at all. The change of form, indeed the actual inter- change I had anticipated between the closely allied species which grow on the two rocky substrata in a state of nature, did not occur at all. If we still take the case of siliceous and calcareous plants, and regard the soil as the source of free inorganic substances which influence the plants, we are forced to assume that greater quantities of one substance will be injurious to one or other of them. The absorbent cells have the capacity of choosing between the substances at their disposal, but this capacity has a definite limit in every species. The cells can absorb as much as they require from a very weak solution of common salt, soda, gypsum, calcium bicarbonate, &., but a concentrated solution of these salts may injure and destroy their structure and function. If it is allowed to act for any length of time on the cells whose function is to absorb. inorganic nutriment, the death of the whole plant will inevitably result. If the Moss which grows on blocks of granite is ‘watered with a saturated solution of gypsum; if the soil into which our Meadow-grasses send their roots is watered with a saturated solution of common salt; or if the humus in which the plants of an upland moor grow is mixed with sodium carbonate or calcium bicarbonate, the plants invariably perish, and the same mineral substances, which in a very weak solution are needful, or at any rate harmless, become poisonous when the solutions are concentrated. The fact that one species of plant prefers this and another that mineral substance (see vol. i. p. 73), however, renders it probable that the injurious effect of materials in large quantity in the soil varies, that a large quantity of Vou. IT. 82 498 DEPENDENCE OF PLANT FORM ON SOIL AND CLIMATE. common salt would be injurious to one species, and an abundance of sodium or potassium salts to another. From the present standpoint of our knowledge concerning the absorption of inorganic materials by plants, therefore, Unger’s classification, especially the expressions silica-demanding and silica-preferring, is no longer suitable, and it would be more to the purpose to speak of plants which are injured by lime, potash, &c. The difference in the vegetation on the closely adjoining limestone and slate mountains met with in so many places in the Al,s, and so well seen in the neigh- bourhood of Kitzbithel, where the climatic influences on the two ranges are identical, can be accounted for most satisfactorily in the following way. Plant-species which demand or prefer a siliceous soil are absent from limestone mountains wherever their roots would be exposed to more free lime than is beneficial; if present they would be weakened, and thus vanquished in the struggle with their fellows, to whom the larger quantity of lime is harmless, and they would eventually perish. These plants flourish luxuriantly, however, on slate mountains, because there the soil does not contain an injurious amount of lime. The absence of species, demand- ing or preferring lime, from slate mountains can be explained in the same way. When seeds are brought thither by the wind from the neighbouring limestone mountains and germination commences, their further development is visibly retarded; they dwindle wherever there is not much lime, and are overgrown and suppressed by the siliceous species which flourish there so luxuriantly. The brown or black mass formed by the decomposition of dead plant residues, known as humus, plays a very important part in the contrasting vegetation on limestone and slate mountains. To obtain a true idea of its significance it must first be pointed out that three distinct stages can be distinguished in the development of a continuous and intricate plant-covering. To the first stage belong the plants which settle down on the bare earth content with a substratum wholly devoid of humus; in the course of time they conquer the-most barren rock, the barest boulders, and the dreariest shifting sands. The species of this group belong chiefly to the Lichens, Mosses, Grasses, Pinks, Crucifers, House-leeks, Saxifrages, and Composites, whose spores, seeds, and fruits are exceptionally well adapted for wind distribution, and can be transferred with ease to the steepest slopes and the most uncompromising crags. The second stage includes plants which require a moderate amount of soil mixed with humus; they establish themselves on the ground pre- pared by the first settlers, wresting it from them and taking possession, and then suppressing and overgrowing them entirely. These plants belong to very different families, whose distribution and establishment are effected in very many ways to be described subsequently. The third stage of development consists of plants for which the abundant humus stored up successively by the plants of the second stage is absolutely indispensable. Bog-moss, Lycopodiums, Sedges, and Heaths form the chief part of this stage. In the course of years the amount of inorganic materials in the soil which supports the plants of the third stage continuously diminishes. Plants which require a large quantity of inorganic salts languish, and are, moreover, INFLUENCE OF THE SUBSTRATUM. 499 overcome by saprophytes which find a suitable habitat there and flourish in abundance, The decayed portions of Saprophytes contain relatively little inorganic material. No trace of lime (in particular) is to be found in their ash. In this way a superficial layer of humus is formed which actually excludes a large number of plants. The next deeper layer may contain a considerable quantity of inorganic salts, but they are valueless to plants rooted in the upper (humus) layer, as they cannot penetrate it. It has been shown by experiment that pure humus possesses the power of holding back materials which are soluble in water. It possesses this property to such an extent that if salt solutions are filtered through a layer of humus the water which escapes below is almost pure. It is therefore impossible for inorganic substances from the deeper layers of the soil, much less from the underlying rock, to reach the surface layer of humus in solution by diffusion; and if some mineral ingredients are not introduced by irrigation or flooding, the upper layer of soil consists of pure humus on which only saprophytic plants can flourish. The formation of such layers of humus occurs much more easily and quickly on slate mountains than on limestone, because in the former the rock and the products of its decomposition retain water much better, and a uniform saturation promotes the development of humus, and also because on slaty soil the second stage of the development of the plant-covering consists of plants which require very few inor- ganic food-substances, and accordingly very few inorganic materials are yielded by the humus, which originates at the cost of the decaying portions of these plants. But a thick stratum of pure humus may also arise in course of time on lime- stone mountains. Only the soil must be uniformly moist in that spot, and neither sand nor mud must be deposited on it. If these conditions are fulfilled a deep humus will gradually spread itself over limestone rocks and débris in the third stage of development, the superficial layer of which will contain no trace of lime, but will afford an excellent soil for silica-loving plants (2.e. for those to which lime is injurious). The isolated occurrence of so-called siliceous or slate- plants on limestone mountains, even in the middle of a patch of plants which are characteristic of a limestone soil, may be naturally explained in this fashion. The water which moistens the rock and soaks the soil has, apart from its mechanical action, the important function of opening up mineral substances and of forming solutions from which the absorbent plant-cells may take their choice. The atmospheric water which penetrates into the earth from above is especially valuable as a solvent on account of the carbonic acid gas it contains. It is immeasurably more valuable to every part of the soil which is riddled by the roots of living plants than the soil-water, so poor in carbonic acid, which collects on impervious strata of the soil and soaks upwards through the superficial layers. The power of the soil to retain water depends mainly on the extent of breaking up undergone by the rock whose disintegration has formed the soil and upon the amount of clay which has arisen from this disintegration. But the amount of humus which in course of time has mixed with the disintegration and the decom- position products of the underlying rock is also an important factor, and thus very 500 DEPENDENCE OF PLANT FORM ON SOIL AND CLIMATE, complex conditions arise which render the estimation of the soil’s capacity for retaining water very difficult. If permeable sandy soil, poor in humus, is deprived of ground water and is dependent for its moisture solely on the atmosphere, the plants growing in it will be retarded in their development if rain and dew are absent for any length of time, and their outward appearance will be altered by this restriction of growth. Annual plants subjected to a lack of moisture in the soil just at the time when their growth should be at its maximum, show best how far these alterations will go. The stem-structures remain short, the foliage-leaves shrink to their smallest extent, and no lateral shoots are developed. Only a few, or perhaps only one, of the flower-buds mature; it is small, opens comparatively very early, and the whole plant has a dwarfed aspect. Annual plants of the Poppy (Papaver Rheas, somniferwm), Pheasant’s Eye (Adonis estwwalis, flammea), Corn- cockle(Agrostemma Githago), Cornflower (Centawrea Cyanus),and common Groundsel (Senecio vulgaris) grown on a dry soil differ from plants grown in the same place, but in a damp year, to such an extent in the size of all their parts that at first sight. they might be mistaken for other species. A clay soil which retains water is less. exposed to danger of too great dryness, but if it is not mixed with humus, and therefore loosened, it has the disadvantage that the water it contains cannot take- up the inorganic foods quickly enough and in sufficient quantity for the require- ments of the plants. This drawback explains the surprising fact that plants grown. on heavy wet clay soils have a dwarfed appearance exactly like plants growing on dry sandy soil. In regions liable to flooding by streams and rivers where not. infrequently sandy and clay soils, in all degrees of porosity and admixed with humus in all possible proportions, are to be met with within a few yards of one another, certain species of plants are to be found growing near together in all imagin- able degrees of size, e.g. Aster Tripolium, Bidens cernua and tripartita, Polygonum lapathifolium, Rumea maritimus, Veronica Anagallis. In places where the seedlings cannot find enough free mineral foods, in spite of the abundant moisture in the soil, the stem rises to some 3-8 cm.; in places which favour the absorption of food, to some 50-80 cm. We will describe only one species, Veronica Anagallis, more in detail. Plants of this species are found with stems 3-5 cm. high and 0°5 mm. thick, with foliage-leaves 6-12 mm. long and 5-6 mm. broad when fully developed. The number of flowers in one inflorescence is about 4-5, the calyx and ripe capsular fruit measure 3 mm. in length. Contrasting with these are plants with stem 30-50 cm. high and 7-8 mm. thick, whose fully-formed leaves are 80 mm. long and 85 mm. broad. There are 40-50 flowers in each inflorescence, and the calyx and ripe capsule measure 4-5 mm. in length. Generally speaking these plants are about ten times as large as the others. If the soils which give rise to: such surprising differences in size are examined it will be noticed that the dwarfed specimens are rooted in a heavy soil devoid of humus, while the large luxuriant. plants flourish in a clay soil which is mixed with plenty of humus, and is therefore very open. Obviously the plants could not obtain from the heavy clay soil what they required for the structure of a vigorous plant, even although the INFLUENCE OF THE MEDIUM. 501 ground was well moistened and warmed; but this they could obtain in abundance from the saturated clay soil containing the humus. It has been already stated that the ground water is less favourable for vegetation than rain and dew on account of its paucity of carbonic acid. But the moistening of the ground by water which wells up from below brings other evils in its train. By this means the soil is over-saturated for a long time, a condition which the roots of most land-plants will not tolerate. When it remains stationary for a long while potassium and sodium salts, and, under certain conditions, humous acids pass into it from the wet earth in quantities anything but advantageous to the plants. Vege- tation, therefore, exhibits a scanty growth in places where the ground water influences the stratum of soil penetrated by roots, and it usually consists of comparatively few species. In low-lying regions, where the ground water rises to the surface, we have the formation of lakes and ponds with variable water-level. Sometimes the plants growing in such places are quite submerged, while at other times their stem and leaves are above water. Land plants do not take kindly to this. Most of them cannot survive very long immersion; they become suffocated, die, and decompose under water in a few days. Only a few species have the remarkable power of growing equally well below or above water, and these are, of course, extremely interesting on account of their form. In accordance with the great contrast presented by the external conditions of life to which these species are temporarily exposed we have a fundamental change both in their outward appearance and in the internal structure of their several organs. In order that the stem and leaves should be held in the best position by the flowing water, the mechanical tissue in submerged varieties of these species is much reduced (see vol. i. pp. 424 and 665). They are also devoid of the contrivances which usually regulate transpiration, since no evaporation occurs under water. Stems grown under water consequently appear limp and flaccid when taken out of it; their leaves, when compared with those growing in the air, are much weaker and more delicate. They have no gloss, but are brighter green in colour, and in the air they collapse and dry up in a very short time. A vertical section through the leaf shows that the number of cells between the upper and lower epidermis is much reduced, and that the cells are shortened in a direction perpendicular to the leaf surface. The foliage-leaves of Veronica Beccabunga, when grown under water, are hardly one-third as thick as those grown in the air, and between the upper and lower epidermis there are only 4-5 layers of short cells, while in corresponding leaves of aérial plants there are 10-12 cell-layers and a distinct division into palisade and spongy parenchyma (see vol, i. p. 279). The shape of the leaf is also much changed under water. In Veronica Beccabunga the difference in aérial and submerged leaves is very slight, consisting only in the shortening of the petiole and in the marginal teeth becoming less marked. In Veronica Anagallis, likewise, the alteration in shape is incon- siderable, but in many others it is very noticeable, and we shall return to it when speaking of the influence of light. 502 DEPENDENCE OF PLANT FORM ON SOIL AND CLIMATE. Plants rooted in the mud of a river-bed, the stems and leaves of which are surrounded by rapidly-flowing water, must possess corresponding strength if they are not to be torn. In comparing two plants of the same species, the one growing in the still water of a deep lake, the other in a rapidly-flowing stream, it will be noticed that the walls of the superficial cells of the latter have become strongly thickened, and that strong bundles of bast-fibres have developed in the cortex of the stem, while in the former only the weakest traces of bast-fibres can be seen. The extraordinary length of stem, petiole, and leaf-blade is also very surprising in plants which grow in rapid water. The Pondweed Potamogeton fluitans, the Rushes Juncus lamprocarpus and swpinus, the Grasses Agrostis stolonifera and Glyceria flwitans are very instructive examples. A plant of the last-named Grass growing on damp soil on the edge of a stream over the water had linear, bluntly-pointed leaves, whose sheaths were on the average 15 cm. long, the blades 23 em. long and 85 mm. broad. After this plant had been sub- merged under rapidly-flowing water in the following year, leaves unfolded, which tapered gradually to a point, with a sheath having a mean length of 47 em., and blades 73 em. long but only 5 mm. broad. The blades produced in running water were three times as long and actually rather narrower than in the air. There was no difference in the number of strands traversing the blade, but they were nearer to one another than in the aérial leaves. The Arrow-head (Sagittaria sagittifolia), which usually grows on the muddy bottom of shallow lakes, raising its leaves above the still water, has gained its name from the likeness of its leaf-blade to an arrow. If it is planted in the bed of a rapid stream so that the leaves during their development are exposed to a vigorous current, the leaf-blade is almost entirely suppressed. What still remains has the form of a spade, but not infrequently all trace of lamina is wanting. The petiole, however, lengthens to 70 cm., and forms a limp, flat, pale-green ribbon 1-2 em. broad, which might easily be mistaken at first sight for the leaf of Vallisneria. Another remarkable change which is effected by submerging growing plants is the non-development of the epidermal structures called hairs, so that the leaves and stems of submerged plants always appear smooth. The suppression of hair- structures is very noticeable in the aquatic variety of Polygonum amphibium. In aérial plants of this species the leaves have short petioles, are lanceolate in shape, and are covered thickly with short hairs, which are rough to the touch; while the aquatic plants have long-stalked, broadly-linear leaves completely smooth on both sides. The humidity of the atmosphere has a marked effect on the form of land plants. Transpiration, which is so deeply concerned in all the vital processes, is carried on very slowly in air which is almost or quite saturated with water- vapour. If plants of a species which usually grows in dry air come into a humid atmosphere, they must be furnished with means for aiding evaporation, On the other hand, plants which grow in dry air must be protected against excessive transpiration. The aids and protective measures were so minutely described INFLUENCE OF TEMPERATURE. 503 in vol i. pp. 284 and 307, that it is needless to repeat them here; but it should be noted that the capacity of plants to construct their tissue as need requires, either for aiding transpiration or for protection against excessive evaporation, is very limited. It must also be pointed out that it is very difficult to distinguish clearly between the direct effect of the humidity of the air and the effects of other influences. Heat and light, as well as the amount of moisture in the soil, are intimately connected with the humidity of the air, but the relations are difficult to estimate. To a certain extent they are interchangeable, and therefore, in most instances, it is impossible to say which extermal influence is the cause of any particular alteration in the tissue concerned in transpiration. For the answer to the chief question, whether it is possible for a change in the conditions of life to cause an alteration of form in the sense of an adaptation, it is really a matter of indifference which influence causes the visible effect. Only here, as in so many other cases, matters are simplified if a certain partiality is permitted in experiments for solving these difficult questions, and if the interwoven influences of soil and climate are treated separately. The effect of heat on growing plants was discussed at vol. i. p. 523. It only remains to say here that the formation of starch and other reserve-foods, as well as the formation of sugar in fruits, is largely connected with heat. Fruits of the same species which ripen under a higher temperature differ greatly in the amount of sugar they contain from those ripening at a lower temperature. It is generally accepted that the size also of the stem, foliage, flowers, and fruit is influenced by heat. The changes which occur when plants in flower, after being for some time in a very warm room are transferred into a cooler room, the other conditions remaining the same, are in particular now recognized. When a large-flowered bulbous plant, eg. the Belladonna Lily (Amaryllis Belladonna), is transferred to a cold greenhouse after opening its first flowers in a warm one, the flowers it here develops at a lower temperature are almost a third smaller than those produced in the warm house. But when the first flowers open in the cold, and the later ones in a warm atmosphere, the former remain small and the latter are larger in size. It is important to emphasize this circumstance in order that the phenomenon here exhibited may not be mistaken for another, in case we should be led to think that the flowers of a plant which first unfold are larger than those which succeed them even when there has not been the slightest alteration in the conditions of light, heat, humidity, &c. It is particularly instructive, when examining the effect of heat on the form of a species, to compare plants grown in water of different temperatures but under conditions otherwise similar. In mountainous districts the springs on the same mountain slope have a different temperature according to their elevation, and yet the same species of plants may be found growing in springs at the foot and high up on the mountain. Let us take as examples plants of Cardamine amara, Myosotis palustris, Pedicularis palustris, and Veronica Beccabunga. These species grow at the foot of the Patscherkofel, near Innsbruck, in the bed 504 DEPENDENCE OF PLANT FORM ON SOIL AND CLIMATE. of streams with a mean temperature of 102 °C., but they also flourish in a stream above the tree-line, at a height of 1921 metres above the sea-level, known as the “Kreuzbrunnen”. Comparing plants of the same species growing under the influence of these different temperatures, the following differences are to be noted :— Plants of Veronica Beccabunga growing in spring water at a temperature of 102° C. were 20-50 em. high, and displayed 4-6 internodes between the bottom in which they were rooted and the level of the first inflorescences. The internodes of the stem were 60-120 mm. long and 5 mm. thick; the leaves springing from the middle of the plant were 40-60 mm. long, 20-25 mm. broad, and each of the flower racemes had 12-16 flowers. Plants growing in the spring water at a temperature of 42° C. were 10-15 em. high with 4-6 internodes between the ground and the level of the first inflorescences. The internodes were 15-30 mm. long and 10-12 mm. thick, and each inflorescence had 12-16 flowers. Cardamine amara, Myosotis palustris, and Pedicularis palustris behaved similarly. There seemed to be no alteration in the form of the leaves and flowers; the corollas assumed a rather deeper tint in the Kreuzbrunnen; Myosotis palustris, which was 20 cm. high at the foot of the Patscherkofel, was 4-5 cm. high in the Kreuz- brunnen, and closely resembled the Eritrichiwm nanum of the Southern Alps in the deep blue of its corollas. Cardamine amara, in the same cold spring, in addition to the shortening of its internodes and diminution of its foliage-leaves, displayed a red colour on the outside of its white petals which was not present in plants at lower levels. The powerful influence of light on the development of plants was discussed at vol. i. p. 371. The question now before us is how far bright and subdued light are able to alter the size, form, and colour of plants. The following is a general review of what has been ascertained in the matter from experiments and direct observa- tion of nature. When plants of a species develop in subdued light they always have higher stems and longer leaves than when grown in bright light, provided, of course, that the conditions of moisture and temperature have been as far as possible identical. This difference is especially noticeable in comparing two plants of a species, one of which has developed in the dim light of a greenhouse in the short days of winter, the other in an unshaded place in the open country during the summer when the light lasts for 16-17 hours every day. The former has a lank thin stem, delicate yellowish-green leaves, and either none of its flowers unfold or else they have a weak appearance and their corollas are pale and flaccid. The illuminated plant has, on the other hand, a compact vigorous stem, dark green leaves, and unfolds a multitude of bright-hued flowers. One only of the large number of experiments which have been performed for the purpose of determining this matter definitely will be mentioned here—one indeed which shows how far the form of the flowers also may be affected. Seeds of a biennial Saxifrage, Samifraga controversa, which were sown in several flower-pots filled with similar soil, produced numerous young plants. A pot with six of these young plants was taken in the autumn into the hot-house; another, likewise containing INFLUENCE OF LIGHT. 505 six young plants, passed the winter under a thick coat of snow in the open. At the beginning of December the six plants in the hot-house sent up from the centre of their small leaf-rosettes slender stalks 10 em. high, whose upper internodes were 22 mm. long and 1 mm. thick. The stem-leaves were yellowish, entire, elongated, 6-7 mm. long and 2 mm. broad; calyx-tube 4 mm. long, 13 mm. broad; calyx-teeth 2 mm. long, 15 mm. broad; petals 35 mm. long, 2 mm. broad; stamens 1 mm. long. It was noted that lateral axes only developed in the axils of the upper stem-leaves, and that the buds of the lateral shoots in the lower leaf-axils atrophied. In the following May strong stems 6 cm. high were sent up from the leaf-rosettes of the plants which had wintered under the deep snow in the open; their upper internodes were 12 mm. long and 2 mm. thick. The stem-leaves were somewhat broadened in front with dentate margin, red in colour, 5 mm. long and 3 mm. broad. The measurements of the parts of the flowers were:—Calyx-tube, 2 mm. long, and 2 mm. broad; calyx-teeth, 15 mm. long, 1 mm. broad; petals, 2°3 mm. long, and 2 mm. broad; stamens, 1 mm. long. From the axils of the stem-leaves flower-bearing shoots developed, which, like the parts of the main stem exposed to the sun, were coloured red. Here then the alterations which certainly are due to the various light influences consist not only in the lengthening and shortening of the stem- and foliage-leaves, but the flowers are correspondingly changed. The petals of the flowers which opened at the New Year when the days were shortest were not only relatively but actually narrower than those which belonged to flowers which opened in the early summer when the days were longest. It has already been stated that the elongation of the leaves and the division of the leaf-lamina into long narrow segments in submerged leaves is associated with the diminution undergone by the light in passing through the water (see vol. i. p. 665). The elongation of submerged leaves is very well seen in the water Star- wort (Callitriche) and Mare’s-tail (Hippuris). In the latter the linear submerged leaves are thirty times as long as they are broad, while the length of the aérial leaves is only 7-9 times their width. In Rorya amphibia the leaves which develop under water are deeply cleft compared with those produced in the air. The aérial leaves of this Crucifer are linear-lanceolate, about ten times as long as broad, with finely toothed margin. Under water the leaves have an elliptical shape, are 2-3 times as long as broad, and the lamina is cleft almost down to the midrib in narrow segments 2-3 cm. long, like a comb or feather. The aérial leaves of the whorled Waterwort (Llatine Alsinastrwm) are grouped in whorls of three. They have an ovate shape, and their margins are finely notched. Lach is traversed by 3-5 veins. The leaves developed under water are divided almost their whole length into 3-4 narrow linear segments, and each whorl looks as if it were composed of twelve leaves. Each segment is smooth round the edge, and traversed only by one central vein. The difference between the aérial and sub- merged leaves of the white-flowered Crowfoots (belonging to the Batrachiwm section of the genus Ranunculus) is even more surprising. Plants of these Crow- 506 DEPENDENCE OF PLANT FORM ON SOIL AND CLIMATE. foots which have developed on muddy but not inundated ground display three- or five-cleft leaves whose segments are light green in colour, shiny, and almost fleshy, and spread out flat. When these plants are grown under water the leaves appear quite different; they become divided into numerous thread-like or hair- shaped segments which have a dark-green colour, and the polished surface has entirely disappeared. The shade afforded by stones, loose earth, undergrowth, and neighbouring bushes and shrubs acts on growing stems, foliage-leaves, and flowers just in the same way as the light-subduing layer of water. In a place near my country house which was formerly used for storing wood and dry twigs, but which had remained unused for a long time, the Creeping Thistle (Cirsiwm arvense) had established itself and formed an intricate growth. The crowded stems attained a height of 80 cm. at the time of flowering and fruit ripening. In the winter of 1885 wood was again stored there in piles 150 cm. high. When, early in the following summer, the new shoots of the Thistle began to spring up they were obliged to content themselves with growing through the dark chinks between the blocks of wood. Many were thus forced to bend and twist, and finally came against some insur- mountable obstacle so that they dwindled in the crevices of the wood-stack without ever reaching the light. Others again which were able to find a fairly straight road through the crevices grew up until they reached the surface of the wood-heap, they then continued to grow 50 cm. higher and unfolded large foliage-leaves on this upper portion. They also developed branches with flower-heads, and from a distance it looked as if a group of Thistles had grown on the top of the wood- stack. The stems had attained a height of 2 metres. The lower internodes were twice as long as usual, the foliage-leaves which sprang from the stalk inside the dark crevices were small, yellowish green, and the buds in their axils did not develop. The Cow-berry (Vaccinium Vitis-Idewa) behaves similarly when its shoots are obliged to grow up to the light through dead tree-trunks. Shoots which force their way in the dark between the bark and the wood of the trunk may reach the height of a metre, while neighbouring ones, springing directly from the soil of the forest are only 15 cm. high. The shoots inside the bark have a reddish colour, and they bear small pale scales instead of dark-green foliage-leaves. From the creeping stems of the White Clover (Lrifoliwm repens) spring erect petioles terminating in three leaflets, and an erect angular stem bearing a flower- head. In sunny places, especially where no neighbouring plants cast a shade, the petioles reach a length of 8 em, and the stem of 10 em. But if dense bushes overshade the Clover, the petiole and stem elongate until the leaflets and capitulum they bear reach the light. Under these conditions petioles 28 cm. long have been found, and stems attaining a height. of 55 em. An extraordinary elongation also occurs in the radical leaves of the Dandelion (Taraxacum officinale) in places where high Grasses and thick bushes shade the moist soil. In the open the leaves reach a length of 20 cm., but in the shade they become twice or three times as long. The lower part of the leaf lengthens most, the free end is comparatively INFLUENCE OF LIGHT. 507 very little altered, and in the central portion the only change is that the lobes and teeth become shorter and less clearly marked. In order to ascertain the effect of covering plants with earth, numerous bulbs of a species of Tulip (Tulipa Gesneriana) were planted at the same depth in one garden bed, and in another some corms of the Spring Crocus (Crocus vernus). Earth was heaped over these bulbs und corms in successive heights of 5, 10, 15, 20, 25, 30, 35, 40, 45, and 50 cm. Naturally the leaf-tips and flower-buds were first. seen in the places where the bulbs were only covered with 5 cm. of soil; in both beds the development was delayed—in the other cases in proportion to the height. of the soil above the bulbs. Some flower-buds of the Crocus appeared above the 20 em. of soil, one of the Tulip above the 30 cm. Numerous leaf-tips of the Crocus appeared above the 35 cm., and a few of the Tulip above the 40 em. of soil. The perianth-tube, the peduncle and the foliage-leaves were almost twice as long as those which had developed under only 5 em. of soil. The flowers were smaller, and unfolded just above the soil; the leaves were narrower and pale yellow in colour as far up as they were covered with the soil. Neither the Crocus nor the Tulip raised their leaves higher than 40 cm. Apparently the reserve-materials stored in the corm and bulb-scales were not sufficient for a further elongation. The stems and leaves of the Crocus and Tulip thus exhibit alterations similar to those observed in the sprouts of Potato-tubers in a dark cellar. We should expect that if moisture and lack of light produce elongation of shoots and various alterations in leaves, a brilliant illumination would have the opposite effect on growing plants. This is indeed the fact. Plants which have been for a year in the shade and have been placed at the beginning of their development. in the following year in the sun display shorter internodes and firmer leaves; they blossom more abundantly, the flowers are of a deeper hue, and in many cases a covering of hairs is formed over the green portions. It is not necessary to mention how far transpiration, which is much more active in the sun than in the shade, is concerned in this; these alterations are certainly produced in the end by sunlight. The effect of brilliant illumination is best seen by comparing plants grown from similar seeds at different elevations, but under identical conditions in other respects. The results obtained in my experimental garden near the summit of the Blaser in the Tyrol, at a height of 2195 m., during the years 1875-1880 illustrate this very fully, and I will briefly recount them here. The seeds of some annual plants were sown in September. The beds were covered with a layer of snow a metre thick throughout the winter. The germination of the seeds took place in the following year soon after the snow melted between the 10th and 25th June. The seedlings therefore developed during the time when the sun was highest and the days longest, and the young plants were exposed to a temperature not lower but rather higher than that enjoyed by plants from similar seeds which began to develop in the experimental beds of the Vienna Botanic Garden in March, when the daylight lasted about 12 hours. The seedlings of several species (e.g. Gilea tricolor, Hyos- cyamus albus, Plantago Psyllium, Silene Gallica, Trifolium incarnatum) were 508 DEPENDENCE OF PLANT FORM ON SOIL AND CLIMATE. killed by the isolated frosts which occurred in each of the six years of the experi- ment, not only in the last week of June, but during July and August; but others — (eg. Agrostemma Githago, Centaurea Cyanus, Ibveris amara, Lepidium satiwum, | Satureja hortensis, Senecio vulgaris, Turgenia latifolia, Veronica polita, Viola arvensis) only underwent a short temporary stoppage of growth from this cause, and opened their flowers at the end of August and beginning of September. In the plants of some species (e.g. Senecio vulgaris, Veronica polita, Viola arvensis) ripe seeds capable of germinating were formed in September. The flowering speci- mens, in comparison with those which had grown during the short days of the spring exposed to numerous night-frosts in the Vienna gardens, displayed extremely shortened internodes. The number of internodes was also lessened, or rather, fewer were developed. For example, where 10 internodes developed in an experi- mental plant in Vienna, in the Alpine garden a corresponding plant would only have 5-6. The same was true of the development of the flowers. While in a plant of Viola arvensis in Vienna the axillary buds of the first six foliage-leaves were suppressed and flowers were not produced until the seventh and eighth leaves, flowers grew from the third and fourth axillary buds in the same species of plant grown in the Alpine experimental garden. The number of flowers on a plant was less, the petals were smaller on the average, and, generally speaking, the annual plants in the Alpine garden had the same appearance as those grown in the plain on dry, sandy soil described on p. 500. It has already been stated on p. 453 that some of the species which are annuals in the valley and on the plain do not die in the autumn in the Alpine garden, but remain alive through the winter and in the following year develop new shoots from the stem. To describe the alterations undergone by biennial species in Alpine regions we will take Libanotis montana (an Umbeilifer) as an example. Its stem in the Alpine garden was 16-24 em. high and developed 5 internodes which were 2-5 cm. long. From the axils of the 5 green stem-leaves sprang lateral shoots which did not branch but terminated in a single umbel, so that the plant only bore 5 umbels altogether. The plants grown from similar seeds in the Vienna Botanic Garden exhibited a stem more than a metre high with 10 internodes each 10-20 em. long. No lateral shoots were produced from the axils of the lower stem-leaves. Those from the axils of the middle and upper leaves were branched and bore several umbels, On an average a plant had about 20 umbels altogether. Over 300 species of perennial plants were grown in the Alpine experimental garden. Only 32 of them blossomed, however. Those whose flowers usually pre- cede the foliage-leaves were in full blossom at the beginning of July, the others, which had to develop a leafy stem before their flowers appeared at the top or in the axils of the leaves of this stem, did not flower until the end of August and beginning of September. Three species of the latter kind will be more particularly treated of here; one species whose stem bears only a single leaf and is terminated by a single flower (Parnassia palustris), one whose stem is beset with decussate leaves and terminates in a loose inflorescence composed of small cymes (Lychnis INFLUENCE OF ELEVATION. 509: Viscaria), and a third whose stem bears alternate leaves and whose flowers are grouped in capitula (Pyrethrum corymboswm). The Grass of Parnassus (Parnassia palustris) from the Alpine garden, when compared with plants grown in the experimental beds of the Vienna Botanic Garden, showed the following measurements:— Vienna Botanic Garden, Experimental Garden on the Blaser. Height of stem ...........00606 20-27 cm. 5-9 cm. Dimensions of leaf............... 3°3 cm. long, 2°4 cm. broad. 10 cm. long, 0°6 cm. broad. Diameter of flower ...........0.. 2°8-3°4 cm, 1:8-2°0 cm. In the Alpine regions, therefore, the plant was only 4 or }{ as high and the leaves only }-} as large as in Vienna, whilst the flowers in the Alpine region had a much smaller diameter than in Vienna. Comparing the hermaphrodite plants of Lychnis Viscaria in the experimental garden of the Blaser with those of the same species at Vienna, we obtain the following :— Vienna Botanic Garden. Experimental Garden on the Blaser. |. Height of the stem, includin : ihe axis of the indbrescenice: ; Se OW ait: 250-20 Dae, Dimensions of lower leaves...... 80 mm. long, 4 mm. broad. | 50 mm. long, 3 mm. broad. Inflorescence............ceeeeeeeeeeeee 80 , , 50 ,, 55 60 ,, » 40 5 53 Reh ecati eoccncarencmls 15 4 oy BB ys 135 Ge ye Be Lamina of petals ............0cceees 10> ss Be | 45 53 fae 68 ,, a Claw of petals.........:cccceecereeees 8 mm. long. 7 mm. long. Plants from the Alpine garden, therefore, when compared with those from the Vienna Botanic Garden, exhibit smaller measurements of stem, leaves, and flowers. The following points were also noted: the number of internodes in plants from the Vienna Garden was 9, of which 5 were on the axis of the inflorescence; each cyme consisted of 3-5 flowers, and the whole inflorescence bore 33-40 flowers. Plants from the Alpine Garden had only 6-7 internodes, of which 3 belonged to the inflorescence; the cymes composing the inflorescence were only occasionally 3-flowered; in most of them only the central flower developed, the two lateral ones being suppressed. The whole inflorescence included only 5-11 flowers. Plants of Pyrethrum corymbosum, from the Alpine Garden, compared with those from the Vienna Botanic Garden (all raised from one batch of seeds) showed the following differences:— Vienna Botanic Garden. Experimental Garden on the Blaser. Height of the stem,............... Dimensions of leaves,............ Diameter of the capitulum,.... Ray-florets,.......:ccsceeseeeeeeenes 950 mm. 170 mm. long, 50 mm. broad. 26 mm. 8 mm. long, 4 mm. broad. 250 mm. 45-50 mm. long, 20 mm. broad. 20 mm. 7 mm. long, 3 mm. broad. 510 DEPENDENCE OF PLANT FORM ON SOIL AND CLIMATE. In this case, again, plants from the Alpine Garden, when compared with those of the Vienna Botanic Garden, had smaller stems, leaves, and flowers. The lobes of the foliage-leaves from the middle of the stem of plants from the Alpine Garden were pinnate, and the pinne were either entire or beset with two small teeth on each side, near the apex. The stem had ten foliage-leaves, the four uppermost of which were much reduced and served as scale-leaves for the lateral shoots arising from their axils. These lateral axes were not branched, and each bore only a single capitulum. There were five capitula altogether. On plants from the Vienna Botanic Garden the lobes of the foliage-leaves from the middle of the stem were more divided, and the pinne were beset on each side near the top with 3-5 teeth. The stem bore 25-27 foliage-leaves, of which the 6-8 upper ones were much reduced in size, and functioned as scale-leaves for the lateral shoots in their axils) These lateral shoots were branched, and each branch terminated in a capitular inflorescence. The total number of capitula was 20-80. From these examples it will be seen that all the parts of plants grown in the Alpine experimental garden were much hindered in their growth. The foliage- and floral-leaves were smaller, the stems shorter, the number of internodes, foliage-leaves, inflorescences, and flowers was diminished. The flowers were relatively nearer the earth, and this was due not only to the diminished number and length of the internodes of the stem, but principally to the fact that the flowers sprang from the axils of the lower stem-leaves. Plants growing in Alpine regions derive a great advantage from these altera- tions, which are chiefly produced during their development by the influence of the long and bright daylight of June, July, and August. If these plants had to produce the same under-structure as their fellows in the Vienna Botanic Garden, 2015 metres lower down, much time would be lost, and the earliest flowers would hardly open before October, at a time when the winter snow is already beginning to fall. But since the number of internodes is restricted, and flowers are developed from the lower stories, it is possible for the plants to blossom at the end of August and beginning of September, and perhaps to ripen their fruits—one of the chief aims of the plant’s existence. To this modification in their mode of development is also due in part the repeatedly-mentioned fact that many alpine plants blossom earlier than those in lower regions. But in order to avoid misunderstanding, it must be expressly stated that in not one of the thirty-two perennial, nor in the biennial and annual species which blossomed in the Alpine experimental garden, was the early flowering hereditary; con- sequently these plants must be carefully distinguished from the so-called asyn- gamic species, which will be spoken of in one of the last chapters in this book. The relation of light to the colouring matters of plants has been repeatedly the subject of careful investigation. All observers agree that the amount of the pigment known as anthocyanin increases and diminishes with the stronger or weaker sunlight enjoyed by the parts of the plant in question, and that the yellow colouring matter of flowers holds a similar relation. Chlorophyll, however, COLOUR AT DIFFERENT ELEVATIONS. 511 is actually destroyed by bright light in plants which are not properly screened, and the green tissue is then blanched and assumes a yellow tint. Since the intensity of the sun’s rays increases with the elevation in mountain districts (see vol. i. p. 525), we should expect that this effect of light would be shown particularly well in plants of high elevation. And this is certainly the case. The flowers of species grown in the Alpine garden on the Blaser at a height of 2195 metres above the sea exhibited, as a rule, brilliant floral tints, and some were decidedly darker than the flowers grown in the Vienna Botanic Garden. Agrostemma Githago, Campanula pusilla, Dianthus inodorus (sylvestris), Gypso- phila repens, Lotus corniculatus, Saponaria ocymoides, Satureja hortensis, Taraxacum officinale, Vicia Cracca, and Vicia sepiwm are good examples of this. Several species, which produced pure white petals in the Vienna gardens, eg. Inbanotis montana, had petals coloured reddish-violet by anthocyanin on their under sides in the Alpine garden. The glumes of all the Grasses which were green, or only just tinged with violet at a low level became 2 dark brownish- violet in the Alpine garden. The abundant formation of anthocyanin in the green tissue of the foliage-leaves and sepals, and in the stem, was particularly apparent. The leaves of the Stonecrops, Sedum acre, album, and sexangulare became purple-red, those of Dracocephalum Ruyschianum and Leucanthemum vulgare violet, those of Lychnis Viscaria and Satureja hortensis a brownish- red, and the foliage-leaves of Bergenia crassifolia and Potentilla Tvroliensis, even in August, had the scarlet-red colour which they usually assume in sunny spots in the valley in late autumn. I must not omit to mention that, according to some of my zoological friends, many animals, especially spiders and snails, which have been transferred from the plains to the mountain-heights, assume a darker tint in alpine regions. A considerable number of plant species, especially those which grow in the valley in shaded or half-shaded places, as, for example, Arabis procurrens, Digitalis ochroleuca, Geum urbanum, Orobus vernus, Valeriana Phu, and V. simplicifolia, Viola cucullata, developed more or less yellowish leaves in the Alpine garden, where they were exposed to the full sunlight. It was mentioned in vol. i. p. 393, that the Flax (Linum usitatissimum), which flourishes in mountain valleys at a height of 1500 metres, where its chlorophyl! is uninjured, nevertheless turns yellow in the Alpine garden at a height of 2195 metres. From this general review of the modifications in plant-form obtained by culture-experiments, a series of important conclusions may naturally be drawn. In the first place we must point out that two kinds of characters are to be observed in plants, those which are the result of certain conditions and properties of soil and climate, and those which appear independently of these external influences. This distinction is so important that we shall illustrate it by two examples. The white Water-lily, Nymphwa alba, develops scale-leaves of ovate or lanceolate shape with no separation into petiole and lamina. The foliage-leaves, however, 512 DEPENDENCE OF PLANT FORM ON SOIL AND CLIMATE. have a rounded petiole and a disc-shaped lamina. These characters are always present whether the seed which produced the plant germinates in a deep lake or in the mud of a marshy meadow. In the marshy meadow the scale-leaves remain short, and the walls of their epidermal cells thicken in a remarkable way; the petioles of the aérial foliage-leaves become about a span long, and, in order to increase their resistance to bending, a strong layer of bast arises, the thickness of these bast-layers amounting to 0°17 mm. The walls of the epidermal cells are thickened, 5-9 layers of collenchymatous cells are formed under the epidermis with walls 007 mm. thick, and the air-spaces in the centre of the leaf- stalk are much narrowed. But if this species of Water-lily grows under water, the scale-leaves elongate into long and flaccid ribbons, and the petioles of the foliage-leaves continue to grow until their blades are raised to the surface of the water. According to its depth they attain a length of 30, 40, 50-100 ecm. Resistance to bending is but little required by the petioles, which are surrounded by water, and the bast is therefore only slightly developed. The strings of bast which traverse the leaf-stalk are only 0°11 mm. thick, the walls of the epidermal cells are only half as thick as in the aérial leaves, only 3-5 layers of collenchyma are developed below the epidermis and the air-spaces in the centre of the leaf-stalk have a diameter of over half a millimetre. These petioles are consequently flexible, and cannot support the leaf-blade if taken out of the water. The general form of the scale- and foliage-leaves, the segmentation of the latter into petiole and blade, the configuration of the blade and the distribution of the bundles in it are all the result of internal forces due to the specific constitution of the protoplasm; but the thickness of the epidermal cells, the strength of the mechanical tissue, and the length of the leaf-stalk, are determined by the depth of the water-covering. The same thing is seen in the flowers; their structure depends upon the specific constitution of the protoplasm, but the size of the petals is determined by the temperature of the water. The Meadow-grass Poa annua has a rapid growth; its haulms and leaf-sheaths are round, the leaf-lamina is traversed by seven strands, the lower branches of the inflorescence are single or paired but never whorled, and the spikelets of the panicle are much compressed and egg-shaped in outline. These characters are unalterable and are observed in Poa annua under all conditions. But when the haulms growing in the gardens in the plain project beyond the short upmost leaf the spikelets become 6-7-flowered, and have a pale green colour. When the plants become perennial in alpine regions the haulms bend towards the ground and remain so short that they do not reach above the highest foliage-leaf; the spikelets develop only 3-4 flowers, and their glumes are dark violet on the surface and brownish-yellow at the edge; thus these modifications are in relation to peculiarities of situation (in the plain and alpine regions) as effect to cause, and are to be ascribed to the influences of heat, light, and moisture, which act in various ways according to the situation. These alterations are always to the advantage of the plant. They make the ARE THESE CHANGES IN FORM PERMANENT? 513 individual more resistant, support and protect its organs, and render it possible for the separate parts to perform their work in spite of the necessarily altered conditions. They seem to have the task of keeping the plant alive under very different vital conditions, of promoting growth and the formation of offshoots and fruit with the smallest possible expenditure, and they may therefore be regarded as adaptations to the particular conditions of soil and climate. The capacity for adaptation is of course founded in the specific constitution of the protoplasm, and is very different in different species. One species may adapt itself by appropriate alterations to the influence of bright light, submersion under water, a dry atmosphere, &c., while another cannot do so. If the protoplasm of the Flax (Linum usitatissimum) could manufacture as much anthocyanin in its green tissue as the Summer Savory (Satureja hortensis) it would blossom and ripen its fruits in alpine regions as this plant does, and would not succumb to the effect of the strong light. If the protoplasm of the Common Bent-grass (Agrostis vulgaris) were able to continue its constructive activity under water it would not perish as soon as it is submerged, but would maintain itself like the stoloniferous species (Agrostis stolonifera) by green stalks and leaves adapted to an aquatic habitat. In short, the adaptability of each species is restricted within definite limits which depend upon the specific constitution of the protoplasm and cannot be overstepped. It is a matter of great import in the history of species whether modifications in form effected by change of soil and climate are transmitted to the descendants, and whether they can be inherited. This of course can only be ascertained by experiments, and by experiments in which all possible sources of error have been eliminated. This last remark is made advisedly, for the sources of error in such experiments are very numerous. I will briefly indicate two which interfered with some experiments I carried out in the years 1863 and 1864. It is not enough to be careful that the seeds sown in the prepared experimental beds are all from the same plant; care must also be taken to see that they are not the result of a hybrid cross-fertilization. Some seeds taken in 1863 from a plant of Dianthus alpinus growing in the Botanic Garden at Innsbruck, and sown in different soil in two experimental beds, produced plants in soil free from lime, which, in their external appearance, agreed with Dianthus deltoides. It seemed as if Dianthus alpinus, a lover of limestone rock, had become transformed into Dianthus deltoides when grown without lime. The seeds of the plant so like Dianthus deltoides were again sown in soil without lime, but the resulting plants no longer resembled this species; they showed themselves to be constant in their characteristics. The whole experiment with Dianthus alpinus was then repeated, but this time the plants on the clay soil without lime did not change, and I was obliged to conclude that. the plant I had regarded as a stage in the transformation of Dianthus alpinus into Dianthus deltoides was a hybrid of these two species. In order to be certain about this a crossing between the two species was effected artificially. From the resulting seed plants were actually grown which were exactly like those I had regarded as transformations, and there was no longer any doubt that some of the Vou. IT. 83 514 THE INFLUENCE OF MUTILATION ON THE FORM OF PLANTS. stigmas of the Dianthus alpinus which had yielded the seeds for the first experi- ment had been pollinated by insects with the pollen of Dianthus deltoides. Mistakes often arise also from the fact that the young stages of many plants are very different from the fully-grown specimens. Young Birches grown from the seeds of Betula verrucosa bear leaves which are simply serrated, thickly covered with hairs, and soft to the touch. They are deceptively like the leaves of adult plants of Betula alba or pubescens. The leaves of the adult Betula verrucosa have quite a different form; they are doubly serrated, smooth, and harsh to the touch. These latter are the only form of leaf described in Botanical books for Betula verrucosa. Anyone sowing the seeds from a grown tree, and watching them grow up, with leaves of a different shape and surface, might easily think an actual fundamental change had occurred, and might be tempted to regard the transforma- tion as the direct effect of a change in external influences. It is perhaps superfluous to state that due regard was paid to these possible sources of error in the later series of cultural experiments, carried out during six years in the Alpine garden on the Blaser (2195 metres), and for comparison in my Villa Marilaun in the high-lying Tyrolese Gschnitzthal (1215 metres), in the Botanic Garden at Innsbruck (569 metres), and in the Botanic Garden of the Vienna University (180 metres); in no instance was any permanent or hereditary modifi-.... . cation in form or colour observed. Seeds of a plant grown in the valley when sown in the Alpine region produced plants which exhibited the modifications described above. They were also mani- fested by the descendants of these plants but only as long as they grew in the same place as their parents. As soon as the seeds formed in the Alpine region were again sown in the beds of the Innsbruck or Vienna Botanic Gardens the plants raised from them immediately resumed the form and colour usual to that position. The modifications of form and colour produced by change of soil and climate are therefore not retained in the descendants; the characteristics which appear as the expression of these changes are not permanent, and the individuals are to be there- fore regarded as varieties, of which Linneus says in his Philosophia Botanica: “Varietates tot sunt, quot differentes plantee ex ejusdem speciei semine sunt pro- ducte. Varietas est Planta mutata a caussa accidentali: Climate, Solo, Calore, Ventis, &e., reducitur itaque in Solo mutato.” THE INFLUENCE OF MUTILATION ON THE FORM OF PLANTS. When Birches and Firs grow up side by side in a wood-clearing, the crowns of the Birches will overtop the Firs in some twenty years’ time, and this will seriously interfere with the growth of the latter. With every blast of wind the whip-like branches of the Birch strike against the upper shoots of the Firs, so that these gradually wither and die off. A lateral branch of a Fir tree altering its direction of growth and replacing the dead leader will, in its turn, soon be scourged to death. The top of the Fir is permanently mutilated, and the injury THE INFLUENCE OF MUTILATION ON THE FORM OF PLANTS. 515 can be recognized years after by the flattened form of the crown, so different from the usual appearance, when the offending Birches have perhaps long dis- appeared. Many other trees wage the same war with one another, the result in each case being the mutilation and alteration of the form of the summit of one of the trees. The Maple, for example, is either put quite hors de combat by the long thorny branches of a neighbouring Gleditschia (Gleditschia triacanthos) or else the crown becomes lop-sided owing to the destruction of the branches on the side facing the Gleditschia. The way in which the appearance of Firs, Larches, Beeches, and Ling is altered by the attacks of ruminants, especially goats, was described in vol. i. p. 445, and we may add here that Pines and Junipers are mutilated in the same manner. The consequence is that lateral branches, which would not otherwise develop, grow out in the following year from the base of the twigs which have been bitten off Apparently no other alteration takes place in these plants. But when huge boughs are broken off close to the ground by storms and the weight of snow, when the tree-trunks of the forest are sacrificed to the wood-cutter’s hatchet, and the stems of seedling trees and shrubs in the meadow to the mower’s scythe, when all the young shoots are frozen by a night’s frost in spring, or when all the leaves are devoured by caterpillars and the branches are left bare as in winter—then the consequences are much more serious. In these cases new shoots make their appearance either from “eyes” in the stem or from the reserve-buds of the branches and twigs, or by buds produced by the roots below the ground. The leaves of these shoots, or suckers, as they are called, differ very much from those of the branches which have been broken, eaten, cut, or frozen off. The leaves from the crown of the Aspen (Populus tremula) are stiff and smooth in their adult condition; the circular blade is borne on a long petiole, and its margin is coarsely notched and undulated. The lateral veins traversing the blade are lost in a network near the edge in which no strong curved ribs occur. The leaves of a sucker from the base of a mutilated stem, or from the root, are soft and thickly covered on both sides with downy hairs; the heart-shaped blade is borne on a short stalk, and the margin is beset with numerous upwardly-directed notched teeth. The lateral veins of the blade merge near the edge of the leaf into a network, in which strong curved ribs are plainly visible. The leaves from the crown of the Oak (Quercus pedunculata) are deeply lobed and furnished with two so-called auricles at the base; those of the suckers are quite entire or very slightly lobed, with no auricles at the base. The leaves of the sucker of the eommon Beech (Fagus sylvatica) are more or less plainly serrated at the edge, while those of the topmost branches of the tree are quite entire. In the Black Mulberry (Morus nigra), and in the Paper Mulberry (Broussonetia papyrifera), the leaves of the sucker have a sinuous margin and are more or less deeply lobed, but those of the tree-top are heart-shaped with notched margins and no lobes. The leaves of the sucker of the Birch (Betula verrucosa) are simply serrated, with velvety hairs; those on the crown of the tree are doubly serrated and 516 THE INFLUENCE OF MUTILATION ON THE FORM OF PLANTS. smooth. The leaves on the suckers of the Round-eared Willow (Salia awrita) are broadly ovate, fairly smooth, and the veins in the blade form a wide-meshed reticulum; the leaves on non-mutilated branches are widened in the upper third, strongly wrinkled, and covered with grey hairs, whilst the reticulum of the veins is narrow-meshed. In Saliva rosmarinifolia, the leaves of the suckers are twice or three times as broad as those of the normal branches, and they are smooth, while those of ordinary branches are covered with silky hairs, and gleam like silver. Hundreds of trees and shrubs might be mentioned in which there is a distinct difference between the foliage of the suckers and of the normal branches of the crown. But these few examples will suffice, and we will only mention the Norway Maple (Acer platanoides), because the difference in the foliage-leaves can be seen from the illustrations in vol. i. The leaves of the summit (see vol. i. fig. 106, p. 416, and fig. 109, p. 419) are borne on long petioles, the blade is 5-7 lobed, and the lobes are short and beset with several pointed, tapering teeth. The leaves of the suckers in this same Norway Maple are short-stalked, the blade is slightly 3-lobed, and each lobe is triangular and without the elongated pointed teeth. They exactly resemble the first foliage-leaves shown in vol. i. p. 9, fig. 1% a ta This is also true of the leaves on the suckers of other woody plants. The shoots - - developed from reserve buds, “eyes”, and the like, repeat to a certain extent the beginning of the leafy stem, so that the phenomenon is only an exhibition of the usual metamorphosis of the foliage-leaves. The difference between the older and younger, 7.¢. lower and upper foliage-leaves, only seems strange because the two kinds of leaf-forms are not usually seen simultaneously on one and the same plant. By the time the crown of a tree has developed, the first (oldest) leaves which adorned the young sapling have long disappeared. Many descriptive Botanists, as a rule, only consider the foliage-leaves of the fully-grown trees and bushes; some of them have hardly ever seen the first leaves of the commonest trees, and when they do happen to come across them they regard them as an extraordinary phenomenon, declare the shoots bearing them to be “bud variations”, and draw bold and bewildering hypotheses from their appearance. This alteration in form, however, has nothing to do with the formation of varieties, nor is it dependent either upon the influence of the soil or upon the effect of climate. Moreover, the form of leaf characteristic of the sucker is not possessed by the secondary shoots which arise from the suckers; these are adorned with the same foliage which occurs on the topmost branches of the tree. Alterations in the scale-leaves as well as in the foliage are brought about by mutilation of the branches. When the upper portions of Willow boughs with their foliage-buds are cut off, leaving the lower portions with the buds of the flower-catkins on them, the small pale scales at the base of the catkins change into green foliage-leaves; the axis bearing these leaves elongates, and the catkins then form the termination of a leafy shoot. Many Willows, e.g. Salia cinerea and S. gramdifolia, by this metamorphosis assume a very unusual appearance. In the following year the branches bearing the flower-catkins, if they are seve eee THE INFLUENCE OF MUTILATION ON THE FORM OF PLANTS. 517 not mutilated afresh, will again put out short catkin-stalks with small pale scales. Mutilation of herbaceous plants is caused by herbivorous animals, viz. insects and mammals, and on a large scale by man when he mows the meadows and cuts the crops and makes other necessary invasions on the natural vegetation in the interests of husbandry. The alterations caused by these mutilations of the foliage-leaf region are in the main the same as in woody plants. From the remaining stumps of the stem lateral shoots arise whose first leaves are like the first leaves of the seedling. Usually they are less divided and have fewer hairs than the leaves on shoots of normal plants, and on this account they have a very different character. In the floral region the effects of mutilation are twofold— first the peduncles or the lateral axes which are terminated by inflorescences elongate, and then the flowers become smaller. For example, when a vigorous stalk of the Ox-eye Daisy (Chrysanthemum Leucanthemum) bearing a capitulum is cut off close to the ground, long lank lateral stems develop from the axils of the lowest remaining leaves, each one ending in a capitulum. The main stem is now seen to be branched at its base, which is never the case in normal plants. If about half the stalk of the common Foxglove is cut off in the spring long flower-racemes will arise from the axils of the leaves just below the cut, but the flowers will be only half as large as those which would have developed on the uncut main stem. The stem of Althwa pallida rises a metre above the ground if its development is not hindered, and forms fascicles of short-stalked flowers in the axils of the upper leaves. If the stem is broken off lateral axes develop from the axils of the remaining leaves, and bear little long-stalked flowers. Particularly good examples are furnished by the annual weeds Delphiniwm Ajacis, Nigella arvensis, Stellera Passerina, and the like, which grow up amongst cereals. Their main stems are broken off when the corn is cut, and they then develop comparatively long branches with small flowers from the remaining stumps. If only single flower- buds, and not the whole inflorescences, are removed from a herbaceous plant whose main stem terminates in a long raceme, so that each flower is cut away in turn from below upwards just before it opens, the rachis of the raceme elongates enormously and flower-buds are developed at its end which would certainly not have unfolded had there been no mutilation. In the Red Foxglove, for example, the rachis of the raceme which has been damaged in this way will grow to twice its ordinary length, and twice as many flowers will be developed. The last and highest flowers in such racemes, however, are only half the size of those which arise on normal racemes. We must now consider certain perennial meadow plants which when mown ‘down are stimulated by the mutilation to develop flower-stalks in the same year, which would, in the normal course of things, not have flowered till the year follow- ing. In Alpine valleys it is a very common thing for the flowers of the spring plants Anemone vernalis, Geraniwm sylvaticum, Gentiana verna, Polygonum Bistorta, Primula elatior and P. farinosa, Trollius Europeus, &e., to appear in 518 ALTERATION OF FORM BY PARASITIC FUNGI. the autumn in meadows which have been mown in the spring. The flowers appear- ing under these circumstances are remarkable for their small size. Their diameter is at least a third smaller than that of the spring flowers. In conclusion we may refer to the gardener’s artifice which has already been described (p. 453) of pro- ducing perennial plants with wocdy stems from an annual Mignonette plant by mutilation, We might also mention the dwarf shrubs and trees produced by combined mutilation and grafting, especially the strange-looking little Ivy trees obtained by grafting a flowering branch of Ivy on an erect stem a span high, and the dwarf Conifers so much in favour with the Japanese. Gardeners and descriptive Botanists have frequently determined and described mutilated plants as other species, hybrids, or varieties. They are neither the one nor the other. The peculiar appearance of the altered members resulting from mutilation is exactly determined beforehand in each species; it is due to the specific constitution of the species, and thus is part of its being. It is not produced by the external influences ‘which lead to the formation of varieties, but is brought about. by inherent necessity, quite independent of the influences of climate and soil. ALTERATION OF FORM BY PARASITIC FUNGL A considerable number of the trees and shrubs of Central and Southern Europe bear bristling, much-branched structures on some of their boughs which, from a distance, look like large birds’ nests or brooms, and which have been popularly termed “witches’ brooms”. They are the outward and visible signs of a disease from which the plants in question suffer, and, as their name testifies, their origin was thought to be connected with witches. Traditionally witches have the power of “wishing” harm to mankind, animals, and plants; and superstitious people, at the sight of these peculiar pathological structures on the trees, may have started the idea that the disease was caused by witches that they might have brooms ready at hand for their midnight ride on the Brocken. Other plant diseases have been ascribed to unusual conditions of weather, especially to long-continued rain or great drought. It is not long since the discovery was made that most of the diseases attacking trees, shrubs, and herbs are caused by Fungi, and that atmos- pheric conditions are only concerned in the matter in so far as they hinder or favour the establishment and development of these parasites. All the Fungi in question are parasites. They penetrate into the tissues of the host-plant and sooner or later cause the death of the affected part, and frequently of the entire host-plant. The living protoplasm in the cells and tissues of the host which is influenced by the parasite undergoes fundamental changes in its com- position. Some of the cells are drained, their living protoplasm being consumed, so to speak, and these cells are obviously marked for destruction. Others are not killed, but changed. The metamorphosis occurs, in the first place, in the consti- tution of the living protoplasts which have not yet completed their development, the change much resembling that known as fermentation in fluid substances TYPES OF FUNGAL GALLS. 519 (cf. vol. i, p. 508). In fermentation the chemical composition of the fluid is altered, its chemical compounds are shaken, decomposed, and split up and new compounds are formed by the action of the living Yeast cells. The same thing happens here in the interior of the living plant in its turgid, meristematic tissue— that is to say, in a group of protoplasts which still have the power of growing at the expense of materials supplied them, of increasing in size, and of multiplying by division. But these cells no longer behave as—in the absence of the parasite— they would have done. Profoundly modified under the influence of the parasite, but yet not killed, these cells, by their continued division, form tissues and organs of new and unusual form; in other words, that part of the host which is invaded but not killed by the parasite will continue to grow and increase in size, and in consequence of the change which its protoplasm has experienced will assume a different outward form. These altered tissue-bodies produced by parasitic Fungi are called gall-structures. They are usually characterized by an excessive growth known as hypertrophy, as well as by their altered shape. The hypertrophy is without doubt caused by a stimulus proceeding from the parasite. We may conclude that the significance of the increased growth lies in the abundant supply of nourishment thus placed at the disposal of the parasite, since the large quantity of food-material brought for the excessive development of the hypertrophied growth connotes a large supply for consumption by the parasite. In many cases, however, the hypertrophied tissue merely forms a wall protecting the host against the further depredations of the intruder. It then contains no nourishment for the use of the parasite, being built up chiefly of corky cells, which the latter cannot consume or destroy. Such a tissue might be compared to the so-called callus which grows up in plants in parts de- prived of epidermis after an injury, or in other wounds, and gradually covers them over with a protective layer. The formation of the gall is often restricted to only a small portion of the afflicted plant; in other cases whole leaves and branches, and sometimes even ex- tensive shoots, become modified in shape. To get a general idea of the four types of hypertrophied growths it will be best to take them one after the other in the order mentioned, commencing with the simplest. The simplest of these galls consist of a few degenerate and metamorphosed cells in the centre of an extensive and unaltered tissue. They are produced chiefly by parasites of the genera Rozella, Synchytriwm, Exobasidiwm, and Gymnosporan- gium. Rozella septigena, one of the Chytridiex, develops swarm-spores which attack the various species of the fungal genus Saprolegnia. They settle on the tubular branches of the Saprolegnia at a place where it was just about to divide and to produce swarm-spores of its own. In consequence of the invasion of the parasite this does not take place, but the tubular cells which would have formed a Saprolegnia-sporangium divide instead into short barrel-shaped cells, each of which becomes a sporangium of Rozella septigena. In addition to this the infected cells develop lateral outpushings which swell up spherically, and each contains a resting- 520 ALTERATION OF FORM BY PARASITIC FUNGI. spore of the parasite. Parasitic species of Synchytrvwm cause a vesicular enlarge- ment of single cells of the epidermis in the leaves of phanerogamic host-plants. The not uncommon species Synchytrium Anemones and S. Taraxact produce only a slight overarching, and the enlargement of the cells is hardly more than four times, often only twice the usual size. But, by the influence of Synchytriwm Myosotidis, hypertrophied epidermal cells rise up from the leaves of the Forget- me-not (Myosotis) in the form of comparatively large, club-shaped, bottle-like, or egg-shaped bladders of golden or reddish yellow colour, and each contains the parasite, or rather its spores. The parts of the leaf attacked by Synchytrium Myosotidis are also much thickened, the palisade cells and the air-containing lacune of the spongy parenchyma (cf. vol. i. p. 279) disappear, and the tissue consists entirely of large similarly-shaped cells which fit close to one another, leaving no spaces between. In the gall caused by Synchytrium pilificwm on Potentilla Tormentilla the much-enlarged cells in which the parasite settles are overgrown by the adjoining hypertrophied cells, some of which rise up in the form of hairs, and the whole new structure resembles a hairy wart. A curious gall is produced by Exobasidium Vaccinit on a sharply-defined portion of the folsage-leaves of the Alpine Rose (Rhododendron hirsutum and ferrugineum). A spherical spongy body rises from a restricted portion of the leaf, usually from the under side of the somewhat projecting midrib, sometimes only as large as a pea, sometimes as big as a cherry, and occasionally even attaining the dimensions of a small apple. It is yellow, but rosy-cheeked like an apple on the side turned to the sunlight, and it reminds one of this fruit by its succulent tissue and sweet taste. Indeed, these galls are sometimes called “Alpine Rose- apples”. Their surface is covered with a bloom which is caused by the numerous spores developed there and does not consist of wax like the bloom on an apple rind. The neck joining the gall to the leaf is not more than 1-2 mm. across, and, what is still more remarkable, except for this sharply-defined place of connection the infected leaf is unaltered. Galls produced by the Gymnosporangia on the leaves of the Mountain Ash, Pear-tree, Rock-medlar, and other Pomez exhibit strange forms. One of them, caused by Gymnosporangiwm conicum, on the foliage of the Rock-medlar (Aronia rotundifolia), is represented in fig. 3577. It resembles a tubercle furnished with horns projecting from the lower surface of the leaf. Microscopic examination shows that the knob consists of the strangely metamorphosed spongy parenchyma of the leaf. The intercellular spaces which normally contain air are quite filled with the mycelial threads, and in the projecting portion of the tubercle, which is very hard and almost cartilaginous, tubes are inserted which terminate blindly below, where the spores of the parasite are developed, whilst above they are open and fringed, thus allowing the spores to escape. These tubes look like horns to the naked eye. Usually several galls occur together on the same leaf. They are conspicuous at some distance on account of their colour. The chlorophyll is destroyed wherever the mycelium of the parasite extends and a reddish-yellow FUNGUS-GALLS ON STEMS. 521 colour takes its place, so that orange spots appear on the surface of the foliage, contrasting vividly with the green of the unaltered portions of the leaf. Galls rising from sharply defined parts of the stem are comparatively rare. One of the most remarkable is produced on the stems of a Laurel (Laurus Canarvensis) by the parasitic Hxobasidium Lauri. When it appears above the bark it looks like an aérial root, but rapidly grows into a branched spongy body 8-12 cm. long similar in appearance to one of the Fungi belonging to the family Clavariez (cf. fig. 195', p. 21). The galls produced by Entyloma Aschersonii and Magnusii on the Composites Helichrysum arenariwm and Gnaphaliwm luteo-album Fig. 357 —Fungus-galls. 1 Gall on the stem of the Juniper (Juniperus communis) produced by Gymnosporangium clavarieforme. 2 Gall on the leaves of Aronia rotundifolia produced by Gy 3p gi i take the form of outgrowths, varying from the size of a pea to that of a walnut, developed from special spots on the root. Whether the spherical tubercles growing on the root-fibres of many Leguminose, especially those of the Bird’s-foot Trefoil (Lotus corniculatus), the Fenugreek (Trigonella fenwm-grecum), Lady’s-Fingers (Anthyllis Vulneraria), Lupin (Lupinus variabilis), and the Liquorice (Glycyrrhiza glabra) are to be regarded as true galls caused by the Bacteria-like organisms invariably to be found in their interior is questionable. According to the most recent investigations they are the outward expression of a case of symbiosis and not of pure parasitism. Gall developments which involve whole roots or rootlets are found on the Alder (Alnus glutinosa), and on the Cabbage (Brassica oleracea). The gall which is produced on Alder roots by Frankia Alni attains the size of a walnut and has a 522 ALTERATION OF FORM BY PARASITIC FUNGI. curious gnarled appearance; all the fibres of the root-branch thicken in a club-like or tuberous manner and become twisted and entangled with one another. The so- called “ Fingers and Toes”, caused by the Myxomycete (Plasmodiophora Brassice), is a gall-like hypertrophy on the root of Brassica oleracea, which not uncommonly grows to the size of a man’s head. Many woody plants have galls which alter the internal structure as well as the outward appearance of large tracts of the stem. The parasites settle in the corti- cal parenchyma, producing hypertrophy there, and afterwards the most varied distortions and alterations in the wood of that region of the stem. The trunk, branch, or twig becomes much swollen or knotted and the cortex rent and torn. Resin or a gummy mucilage sometimes runs out of the rifts in the gall. As such a parasite exercises its metamorphosing faculty for several years, the canker (as it may be termed) increases in size continually. Sporangia of varied form and colour appear annually on the affected places, and again disappear when they have shed their spores. The part of the stem or branch above the cankerous cushion dwindles and dies off sooner or later. It rarely happens that the tree or shrub is able to rid itself of the parasite. Occasionally a growth of wood and cork from the adjoining healthy part walls in the cankerous spot so that the parasite is destroyed. The gall produced by Gymmnosporangium clavarieforme on the trunks and branches of the common Juniper (Juniperus communis) is an example of this form (see fig. 357"). From the hypertrophy there project in the early spring golden- yellow tongues (shown in the figure) consisting of masses of spores embedded in mucilage. Other similar growths are produced on species of Juniper by Gymno- sporangvum conicum, G. Sabine, and G. tremelloides, but it would take too long to describe their differences in detail. It is important to mention, however, that each of these parasites has two stages of development, living on different hosts, the hypertrophies as well as the associated spore-producing organs of the parasite being different in the two cases. The “ Aicidium stage” produces carti- laginous swellings (see p. 520) in definite spots on the foliage of various Pomer (Aronia, Crategus, Pyrus, Sorbus), the “ Teleutospore stage” thickenings and tuberous outgrowths on the trunks of Junipers (Juniperus communis, excelsa, Sabina), and these parasites can travel from one host to the other in turn. (The two stages on different hosts are shown in fig. 357; these are not of the same fungus, but of nearly allied ones, and illustrate the point mentioned.) The parasite Peziza Willkommii attacks the trunks and branches of the Larch (Larix Europea), and produces the well-known Larch-disease or “Larch-canker”. The parasite having gained access at some point on the stem or branch first pene- trates the cortical parenchyma, and affects the cambium so as to prevent the further development of wood in that place. The development of the wood on the opposite side of the stem, ie. the formation of annual rings, may proceed for several years, and in this way the attacked spot on the trunk takes the form of a depression, which is rendered the more conspicuous should the wood and cortex surrounding the parasite have undergone a greater thickening than usual. In ENTIRE LEAVES AFFECTED BY FUNGI. 523 time the patch becomes a sunken, blistered hole from which resin flows; and every year the fructifications appear above the cortex in the form of numerous little cup-like structures which are white outside and scarlet-red in the concavity. As the disease progresses the infected patch gradually spreads, and infected trunks and branches can be easily distinguished at a distance. Towards the end of summer the needles on the twigs above the canker turn yellow, while those on the healthy branches are still a beautiful green. This premature discoloration is a sure sign of the speedy death of the whole bough. A similar canker is produced on the Fig. 358.—Various Galls. 1G@all on the bract-scales of the pistillate flowers of the Gray Alder (Alnus incana) produced by Exoascus Alni-incane. 2 Inflorescence of Valerianella carinata. % The same inflorescence with galls produced by a gall-mite, +4 Leaf rosette of the House-leek (Sempervivwm hirtum): 5 Leaf rosette of the same plant which has been attacked by the fungus Endo- phyllum Sempervivi and has become hypertrophied. Silver Fir (Abies pectinata) by Acidiwm elatinwm, but instead of being only on one side of the branch, as in the Larch, it forms a uniform swelling all round it. Cankers of this kind are produced by a Bacterial organism (Bacillus amylovorus) on fruit-trees (Apple, Pear, &c.), and on various trees belonging to the Amentifere (Beeches, Hornbeams, Oaks, &c.) by the Fungus Nectria ditissima. When whole leaves undergo hypertrophy of the kind we have particularly remarkable changes of form. For example, the normal leaves forming the rosettes of the House-leek (Sempervivum hirtwm; see fig. 358*) are broadly obovate in form, being little more than twice as long as they are broad. The leaves of the same plant after they have been attacked by the parasitic Endophyllum Semper- 524 ALTERATION OF FORM BY PARASITIC FUNGI. wivi (see fig. 358°) are seven times as long as broad and linear in shape. They stand erect, and are of a much paler colour than the healthy leaves. The Wood Anemone (Anemone nemorosa) affords another example (see fig. 259, p. 229). It spreads by creeping stems under the surface of the ground, and forms small colonies in light thickets and in meadows. The plants consist partly of flowering lateral shoots, and partly of foliage-leaves, which emerge above the ground from the creeping underground stem. In normal leaves the erect petioles are all the same length, and the leaflets are extended at about the same level. But when the Aicidium stage of Puccinia fusca has settled on them this becomes altered. The blades of the infected leaves tower over their healthy neighbours in consequence of the elongation of their petioles, whilst their leaflets are smaller and less divided. The length of the petiole in normal leaves is some 12-13 em., in hypertrophied leaves 15-18 cm.; but the size of the altered segments, compared with those of normal leaves, is as 5:7. Similar changes are observed in leaves of Soldanella alpina when attacked by Puccinia Soldanelle. The petioles of the infected leaves are 2-4 times as long as the normal ones, the blade is smaller and hollowed like a spoon instead of being flat, and the colour is an ochreous yellow instead of a dark green. The same alterations in the length of the petiole, and, in the size and colouring of the- leaf-lamina, are produced in the leaves of Alchemilla vulgaris by Uromyces Alchemille and in those of Phytewma orbi- culare by Uromyces Phytewmatum. To this class belongs also the so-called “curl” disease of Peach and Almond trees, produced by Exoascus deformans, and rendered conspicuous by the considerable enlargement, undulation, and bladder- “like expansion of the infected leaf-surface, which acquires generally a very brilliant coloration. Floral-leaves are comparatively seldom metamorphosed by Fungal parasites. In the Alder (Alnus glutinosa and incana) the bracts of the pistillate flowers are changed by Exoascus Alni-incane (=E. amentorum) into elongated purple-red spatulate lobes much twisted and bent (see fig. 3581); Peronospora violacea some- times causes the stamens to change into petal-like structures in the flowers of Knautia arvensis, so that they then seem to be “double”; Ustilago Maydis causes a growth of tissue in the pistillate flowers of the Maize, the result being that instead of grains irregular cushion-like structures 7 cm. in diameter are produced. Taphrina awrea, which settles on the pistillate flowers of Poplar (Populus alba and tremula) causes the ovaries to form golden-yellow capsules more than twice the usual size. The galls produced by Exoaseus Prwni on the ovaries of wild Plum, Bullace, Sloe, and Bird Cherry (Prunus domestica, insititia, spinosa, Padus) belong also to this class. The tissue of the ovary increases in size, but not in the same way as in fruit forma- tion. The resulting body is flattened on two sides, brittle and yellow; the seed inside is abortive, and a hollow space is left in its stead. The gall produced from the ovary of Prunus domestica has the form of a rather curved pocket, which looks as if it had been powdered outside with flour at the time the spores ripen. These hypertrophies, which are popularly termed “pocket-plums”, “ bladder-plums”, ENTIRE SHOOTS AFFECTED BY FUNGI, 525: &e., fall off the trees at the end of May. They are eaten in many districts, but have an insipid, sweetish taste. Galls consisting of whole shoots, both the stem and its leaves being altered by the parasite, are found principally on trees and shrubs, and only rarely on herbaceous plants. Examples of the latter, however, are furnished by the metamorphosed shoots of the Shepherd’s Purse (Capsella Bursa-pastoris) produced by Cystopus candidus and Peronospora parasitica. Here the leaves, especially the floral-leaves, as well as the ground-tissue of the stem undergo pronounced hypertrophy. The petals, which measure only 2 mm. in length in a healthy plant, may become even 15 mm. long; the sepals also elongate, become fleshy and brittle, and are distorted and crumpled in all manner of ways. Only six stamens are developed in normal flowers, but in hypertrophied specimens there are often eight. The metamorphosis produced by Uromyces Pisi in one of the Spurges, Huphorbia Cyparissias, is even more remarkable. The stem elongates far beyond its usual dimensions, and the leaves, which are crowded together on normal shoots, are thus separated by con- siderable intervals. The distance between two adjoining successive leaves in the healthy Huphorbia Cyparissias is only 0°5 mm., but in the hypertrophied specimens. it becomes 2-3 mm. Infected shoots on an average are twice as high as healthy ones. The foliage-leaves, which are thin, flexible, linear, and twelve times as long as they are broad in the healthy plant, become, in the infected specimens, thick, brittle, elliptical, and only 2-3 times as long as they are broad. The bluish-green colour of the normal plant is changed into a yellow-ochre tint, and this contributes not a little to the odd appearance of the plant. Affected plants are not uncommon in Switzerland; a locality in which this disease has been very prevalent in recent years being Saas-Fée in the Saas-thal. The metamorphoses produced on the shoots of Periwinkles (Vinca herbacea, major, and minor) by the Uredospore-stage of Puccinium Vinee and on shoots of Cirsiwm arvense by the Teleutospore-stage of Puccinium suaveolens are very like those of the Ewphorbia just mentioned, since the stem becomes much elongated and the leaves shorter, broader, yellow, and brittle. When flowers are developed on these affected shoots, they are more or less abortive and sickly, and no fruits or fertile seeds arise therefrom. Frequently the shoots ‘blossom prematurely. For example, we can at once detect by its elongated rosette- leaves when Primula Clusiana and minima are infected by Uromyces Primule imtegrifolic, and it may be observed when this is the case that the shoots do not wait until the next spring to develop the flowers laid down in the summer, as usual, but open them in the autumn of the same year instead. The Cowberry (Vacciniwm Vitis-Idwa) is especially worthy of notice among low woody plants, because two kinds of parasite attack its shoots. Melampsora Geppertiana, in the Teleutospore-stage, causes a marked, gouty thickening in the cortical parenchyma, which is converted into a spongy tissue; at first it is flesh- coloured, but soon assumes a chestnut-brown tint. The stems elongate very much and grow vertically upwards; and when several of them close together are thus attacked they present a besom-like appearance. The foliage-leaves are much 526 ALTERATION OF FORM BY PARASITIC FUNGI. farther apart than in the healthy plant on account of this stretching of the stem. The lower leaves of the shoot are transformed into small fringed scales, and the upper ones are so much shortened that their outline becomes almost circular. The second parasite to which the Cowberry shoot is subject is Exobasidiwm Vaccinia (a near ally of the already mentioned Hxobasidiwm Lauri, p. 521). The stem becomes pale rose-red colour, and rather thickened and spongy, but it does not elongate much more than usual; the leaves become blistered and curiously convex on the under surface. The substance of the infected leaves becomes brittle and loses its chlorophyll. A red tint appears in place of the green, especially on the upper surface of the leaf, whilst the lower surface, on which the spores develop, looks as if it had been dusted over with flour. Usually the buds develop prematurely on these shoots, 7.¢. the buds which, under ordinary circumstances, would not develop until the next year push out and form new shoots shortly after they have been laid down. The axes of these shoots, however, remain short; their leaves are closely crowded, red in colour, and sessile. From a distance the premature shoots look like large double red flowers inserted in the dark green of the non-infected Cowberry bush. The shoots which develop prematurely on the shrubs of the Bog Whortleberry (Vaccinium uliginosum) by the action of Exobasidium Vaccinti are often met with in alpine regions, and are even more noticeable on account of their fiery-red ALTERATION OF FORM BY GALL-PRODUCING INSECTS. 527 colour. The Bearberry (Arctostaphylos Uva-wrsi), Ledum palustre, and the Marsh Andromeda (Andromeda polifolia) are subject to similar metamorphoses at the hands of Haobasidiwm Vaccinit, so that Vaccinium Vitis-Idea may be regarded as typical of them. When the shoots of the larger shrubs or trees are metamorphosed by parasitic Fungi attacking their branches, we have the formation of the structures popularly termed Witches’ brooms, which were mentioned at the beginning of this chapter. The stimulus necessary for their formation is afforded in different plants by different parasites; on Barberry bushes (Berberis vulgaris) by dicidiwm Magel- henicum (to be distinguished from the common . berberidis), on the Gray Alder (Alnus incana) by Exoascus epiphyllus, on the Hornbeam (Carpinus Betulus) by Exoascus Carpini, on the Bullace (Prunus insititia) by Exoascus imsititie, on other species of the genus Prunus by Exoascus Cerasi, on the Birch (Betula verrucosa) by Eaoascus turgidus, on the Weymouth Pine (Pinus Strobus) by Peridermiwm Strobi, and on the Silver Fir (Abies pectinata) by Hediwm elatinum. Witches’ brooms also occur on the Mastic tree (Pistacia Lentiscus), and on Beeches, Pines, Larches, Spruce Firs, &c., although hitherto we have not been able to ascertain definitely what parasitic Fungi are the cause in these cases. The Witches’ broom of the Silver Fir has been selected and figured (see fig. 359) as a type of these peculiar structures. It always grows on one of the horizontally projecting lateral branches of the Fir, and raises its erect or curved twigs from the upper side, resembling, as it were, an epiphyte growing on the bark of the horizontal bough. The twigs are grouped in whorls and not in two rows, as usually happens in the lateral shoots of the Silver Fir. They are all shortened and thickened, and remarkably soft and pliable, because the cortical parenchyma has become spongy and the wood is only slightly developed. The buds, which in healthy tissue are egg-shaped, are almost spherical here. As in other instances of hypertrophied plant-members, we have a precocious development, a so-called “prolepsis”, in these Witches’ brooms. The buds swell earlier and unfold earlier than those of healthy twigs. The leaves remain short, yellow, somewhat crumpled, and fall off when a year old, while those of normal twigs are long, linear, straight, dark green on the upper side, and remain in position from 6-8 years. The growth of the twig is restricted; it dies off in a few years, and then, inserted on the dark green branches of the Silver Fir, remain the dry, bristling brooms, whose appearance has stimulated the imagination of the peasantry and given rise to the superstitions alluded to at the beginning of this chapter. ALTERATION OF FORM BY GALL-PRODUCING INSECTS. “Certain members of the Arachnoidea, Diptera, and Hymenoptera, which attack and penetrate the tissues of living plants and incite the formation of peculiar excrescences, are known as gall-mites, gall-gnats, and gall-wasps. The 528 ALTERATION OF FORM BY GALL-PRODUCING INSECTS. growths, like small rosy-cheeked apples, which occur on the foliage of Oaks, popularly known as “oak-apples”, are amongst the best known. The terms “gall” and “gall-apple” were used by writers in the sixteenth century, and (like the Old English word galle, the French gaille, and the Italian galla) are derived from the Latin word galla, used for these outgrowths by Pliny in his Natural History. The sixteenth-century writers distinguish between “gall-nuts” and “gall-apples”, meaning by the former the small hard outgrowths on the leaves of Beech-trees. Afterwards the word gall was used for all the outgrowths produced by animals on green living plants. More than that—the hypertrophies described in the preceding chapter, produced in green host-plants by the various families of Fungi, are also included under the term. It has been proposed recently to substitute the word cecidium for gall, and to distinguish the excrescences as myco-cecidia, nemato-cecidia, phyto-cecidia, diptero-cecidia, &c., according as they owe their origin to Fungi, Thread-worms (Nematodes), Gall-mites (Phytoptus), Gnats (Diptera), &c. A systematic classification of this sort, on the lines of the classification of animals, might be of use to Zoologists, but to the Botanist its value is only secondary. He must, as in other similar cases, keep to morphology as the primary ground of classification, and has to arrange the structures according to their agreement in development. Moreover, in a general review, it is necessary to consider whether a whole group of plant-organs or one alone undergoes metamor- phosis; and the starting-point of the outgrowth must also be ascertained; ie. whether it is the foliage-leaves, floral-leaves, stems, or root-structures, &¢., which are the head-quarters of the excrescence. When the gall originating as the nest or temporary habitation of a single animal or colony of animals is limited to a single plant organ it is said to be simple; if, on the other hand, several plant organs are concerned in its production it is said to be compound. Simple galls may, for convenience of description, be divided into (1) Felt- galls, (2) Mamtle-galls, and (8) Solid galls. The Felt-galls are chiefly due to hypertrophied epidermal cells growing out into hairy coverings of various sorts and shapes; Mantle and Solid galls, however, are rather more complicated. In both cases insects are present in swellings of various descriptions, but there is this essential distinction: The Mantle-gall is a hollow structure which, though it may arise in various ways and assume a multiplicity of forms, always has a portion of the surface of the affected organ for its lining—in other words, it is a chamber formed by hypertrophied growth around the place occupied by the insect. In the Solid gall, on the other hand, some spot is pierced by an insect and the eggs deposited im the tissues (not on the surface), the punctured spot forms a swelling with the larva inside, but the lining of the chamber is in no sense a portion or development of the original surface of the organ affected. Again, whilst in most mantle-galls the cavity of the gall is in Open communication with the outside, and the insect can escape by this aperture (though this is not invariably the case), in the solid gall there is not such opening, and the insect FELT-GALLS. 529 has to bore its way out. Needless to say, of both these types there are numerous modifications, but they fall into the two classes (of mantle and solid galls) according to their mode of development. The majority of felt-galls are produced by gall-mites. They form cottony or felted growths on limited and sharply defined areas of green leaves and stems, the surface of which is otherwise smooth, or possesses but few hairs. Some- times they have the form of small tufts, bands, or stripes, sometimes of large spots with irregular contour. In most instances the felt is situated on the under side of the foliage-leaf, and the gall-mite usually prefers the projecting veins to the green surface. In the Lime, Alder, Hornbeam, and Horse-Chestnut, the mites usually establish themselves in the angles formed by the lateral strands where they arise from the midrib, the projecting veins forming the framework for the felted hairs. In the Bramble (Rubus) and the Burnet (Poteriwm) it sometimes happens that the felt is continued down from the lamina to the leaf-stalk, and occasionally the green cortex of the succulent twig is covered with felted bands and spots. In some Brambles and Cinquefoils the sepals become furred by the action of gall-mites, the usual consequence being that the outline also becomes distorted. A swelling or slight hollowing of the green leaf-tissue very frequently accompanies the formation of felted galls, in which case the hairy covering is only visible on the concave side whilst the other remains smooth. This is most remarkable in the foliage of the Avens (Gewm), Vine (V2tes), and Walnut-tree (Juglans), where a dozen white or brown-felted pit-like depressions are sometimes to be seen on the under side of a single leaf. The colour of the felted hairs is white in the leaves of Beeches, Limes, Bird Cherry, Brambles, Cinquefoils and Burnets, green in the common Maple, yellow in the Spindle-tree (Zuonymus verrucosus), sulphur-yellow in Alnus orientalis and Black Poplar (Populus nigra), carmine red at first and then violet in Alnus viridis and in the Birches (Betula alba, carpatica, &c.), and brown in the Avens (Geum macrophyllum), Horse- Chestnut (Zisculus Hippocastanwm), and in the Aspen (Populus tremula). The felted galls which are light in their young stages usually take on a brown tint afterwards. Microscopic investigation has shown that in the formation of felted galls, the epidermal cells, originally tabular in shape and closely fitting, swell out and become transformed into bent and twisted tubes generally shaped like a club or retort, the stimulus being afforded by a minute gall-mite (Phytoptus). These cells look like short hairs to the naked eye, and as they stand side by side in large numbers the covering has a velvety or felted appearance. The mites which produce the felt, deposit their eggs in the juicy hair-shaped cells, and their young live on the materials contained in them. It should be mentioned that formerly these velvety and felted coverings were regarded as Fungi, and were described as distinct genera under the names Hrinewm and Phyllerium (e.g. the gall known as Erinewm quercinum on the leaves of Quereus Cerris). To this group belongs also the gall occurring on the Wood Meadow Grass (Poa nemoralis) con- sisting of cells which resemble root-hairs, which is produced by the gnat Hormo- Vo. IL. 84 530 ALTERATION OF FORM BY GALL-PRODUCING INSECTS. myia Pow. The hair-shaped cells are epidermal, and spring from the stem above the nodes; they break through the leaf-sheath which proceeds from the adjacent node, and are arranged in two groups, which grow in opposite directions, so as to wrap round the stem from the two sides. The whole hairy mass looks as if it had been parted into two. At first the hairs are white; later they become light brown, and when the gall is fully developed they have the form of brown felted strands, wound round the stems and firmly inclosing the larva of the gnat in question. A large number of simple galls are grouped together under the name of Mantle- galls. The insects which give rise to them spend their lives on the surface of the leaves, where they multiply and attach their eggs to the epidermis. A growth is excited in certain layers of the cell-tissue by the stimulus which the animals exercise on their place of settlement. Cavities are thus formed which serve as dwellings for the animals and their brood, and which surround them like a pro- tecting mantle. Mantle-galls may be divided according to their structure into scroll-, pocket-, and covering-galls. Scroll-galls are caused by gall-mites, leaf-lice, tree-hoppers, and flies, and usually occur on the blades, rarely on the petioles of the leaves. The surface inhabited by these animals, which, in the ordinary course of things would have spread out flatly, grows more luxuriantly on one side than on the other, and the result is the formation of a scroll, ¢.e. of a chamber in which the animals are hidden. It is always the side on which the animals live which becomes concave, and the leaf is usually curled up lengthwise. In the Alpine Rose (Rhododendron), Crane’s-bill (Geranium sanguinewm), and Orache (Atriplex hastata, oblongifolia, &c.), it is the upper side of the leaf which is tenanted by the insects, and is therefore the one to roll up; it is the lower side, however, in the Buckthorn (Rhamnus cathartica) and the non-climbing species of Honeysuckle (Lonicera alpigena, &.). In many instances the whole leaf-lamina is rolled up, but more frequently the alteration is restricted to the edge of the leaf when the margin appears to be bordered with a swollen hollow cushion often corrugated or undulating. In the Alpine Rose (Rhododendron ferruginewm and hirsutwm) both halves of the leaf-blade are rolled round (see figs. 3602 and 360°), but usually the rolling is so slight that the gall has the form of a boat or hollow trough. Some- times an alteration in the shape of the leaf accompanies the rolling. For example, the foliage of the Abele (Populus alba) on which Pachypappa vesicalis establishes itself when the leaves are very young, exhibits in addition to the rolling a deep hollowing of the blade. Instead of the short blunt lobes, long pointed segments are formed, which stand side by side when they are rolled up, and cross over one another in many ways so that the mantle-gall on the hollow side is shut in by a veritable lattice-work. The parts of the tissue brought into contact by the rolling do not fuse together, and therefore the cavity in which the gall-producing insects live is always in open communication with the exterior. In most cases the tissues concerned are thickened, brittle, more or less devoid of chlorophyll, and yellow in colour. Not infrequently a red pigment is formed in them, so that the outside of the gall has a yellowish-red colour. The scroll-gall produced by the hemipterous FORMS OF MANTLE-GALLS. 531 Trioza Rkamni on the margin of Buckthorn (Rhamnus cathartica) leaves is very hard and thickened like cartilage. In many plants the epidermal cells lining the gall elongate into hairy structures, as in the felt-galls previously described. Their juicy contents are used as food by the young gall-mites. This is the case, for . example, in the Alpine Rose (Rhododendron ferrugineum, ef. fig. 360°). Pocket- galls are closely allied to the scroll-like forms. The tissue of the leaf-lamina or Fig. 360.—Galls. ! Covering-galls on the petiole of the Black Poplar (Populus nigra) produced by Pemphigus spirotheca. 2 Scroll-galls on the leaves of an Alpine Rose (Rhododendron ferruginewm) produced by gall-mites. % Transverse section of one of these galls. #and 5 Bud-galls on the branchlets of the Wild Thyme (Thymus Serpyllum) produced by gall-mites. © Blister-like galls on the leaf of the Red Currant (Ribes rubrum) produced by Myzus ribis. '7 Part of the leaf seen from below. 8 Vertical section of a portion of this gall. 9 Solid gall on the leaf of the Gray Willow (Salia incana) produced by Nematus pedunculi, 10 The same gall cut open. 1 Part of the wall of this gall in vertical section. 1, 2, 4, 6, and 9 natural size; Sand 6x 4; 8and7 x 8; 8 and 11 x 50. petiole and sometimes that of the cortex in young twigs is subjected to a stimulus where the animals (gall-mites, leaf-lice, diptera) settle, with the result that a hollow protuberance arises whose excavated cavity serves as a temporary dwelling for the insects. The protuberances exhibit a great variety of form and shape, and they differ considerably in their internal structure. The following are the most notice- able forms. First, the plaited galls. They form deep, plaited, sometimes twisted channels in the leaf-tissue which open on the upper side by a narrow hole, and 532 ALTERATION OF FORM BY GALL-PRODUCING INSECTS. project like weals on the lower surface of the leaf. The growing tissue which forms the floor of the channel is yellow and often lined with short hairs. The channel usually follows the course of the larger veins of the lamina, and some- times actually traverses one. Plaited galls are produced by gall-mites. The best known are those on the foliage of Carpinus Betulus, Clematis Flammula and C. recta, and Ribes alpinum. Wrinkled galls come next to the plaited form. The protuberance is here limited to the green tissue shut in by strong rib-like projecting veins, and is only shallow; the upper side of the leaf has bulgings and protuberances and the lower pits and cavities. The protuberances are always developed in numbers close together, so that the leaf looks very much wrinkled in that region. Examples of this form are furnished by the wrinkled galls on the Elm (Ulmus campestris; see fig. 361 *) produced by the leaf- louse Schizoneura Ulmi, and on the Red Currant (Ribes rubrum; see figs. 360 ® 7 8) by another leaf-louse, Myzus ribis. In the latter several wrinkles are usually united into large blister-like protuberances, red on the outside, and covered with jointed cellular structures bearing glands which look to the naked eye like short hairs. This form, though resembling certain felted galls, is distinguished from them by the different form of the hairs arising in consequence of the stimulation. In the Mouse-ear Hawkweed (Hieractwm Pilosella) leaf-fleas (Psyllodes) produce minute protuberances with narrow mouths, which stand out from the lower side turned towards the ground like small warts, and when they occur close together give a corrugated appearance to the leaf. Hollow protuberances of this sort arising upon restricted areas of the leaf-surface, and growing very actively, give rise to bag or sack-like structures attached by a very narrow neck. From their resemblance to a head such galls are sometimes termed capitate galls (Cephalonion). In others, where the outgrowth is fairly thick-walled and in form horn-like, the designation horn gall (Ceratonion) has been given. Between these forms numerous intermediate forms exist which may be compared to pockets, bags, nails, &e. Many of these galls project from both the upper and lower side of the leaf, as though a nail had been driven through it—hence the last-mentioned name. The capitate-gall of the Sloe (Prunus spinosa), caused by a gall-mite, projects almost as much from the under as from the upper side, whilst the similar gall on the foliage of the Bird Cherry (Prunus Padus) rises on the upper side as a long pocket, but below only projects like a small wart. Many capitate and horn-like galls are developed only on one side, and here again there is a very great variety. When the protuberances are due to mites the cavity always opens on the lower side of the leaf. Both the inner wall and mouth of the cavity are covered with hairs, and sometimes the aperture seems to be actually plugged up by them. In the bag-shaped protuberances produced by the leaf-louse Tetraneura Ulmni on Elm leaves, a relatively large slit is formed just at the narrow part of the bag at the moment when the insects leave the cavity (see fig. 361°), The external surface of the protuberances caused by mites on the foliage of Alders (Alnus), Maples (Acer), and Limes (Tilia) is smooth, in those of the Bird Cherry (Prunus Padus), FORMS OF MANTLE-GALLS. 533 and Wayfaring Tree (Viburnum Lantana) ciliated, whilst in the inflated galls of the Elm caused by the white woolly leaf-louse (Schizoneura lanuginosa), it is covered with fine hairs like velvet. The capitate galls on the foliage of Maples, Alders, and Limes, of the Guelder-rose and Strawberry, are scattered abundantly over the whole lamina; in the Sloe they stand out chiefly from the margin of the leaf, and in Elms they occur singly or in groups on its central portions. The size of these galls depends upon their distribution. Those which rise in hundreds from the Fig. 361.—Galls. 18 Solid galls on a Rose-leaf; 1 of Rhodites Rose, 2 of Rhodites Eglanteric, 3 of Rhodites spinosissime. 4 Wrinkled galls on an Elm-leaf (Ulmus campestris) produced by Schizoneura Ulmi. 5 Purse galls on the same leaf, produced by Tetraneura Ulmi. 6 Covering gall on the same leaf, produced by Tetraneura alba. 7 Solid galls on the leafof the Purple Willow (Salix purpurea), produced by Nematus gallarum. 8 Solid galls on the leaves of the same Willow, produced by Nematus vesicator. same lamina have a diameter of 1-3 mm., while those which occur singly or in small groups, often attain a diameter of 2-3 em. Contrasting with these embossed or pocket-galls are the covering galls, forming a third type of mantle-gall. In these, as in the embossed forms, the insects pro- ducing the galls live in their cavities, but the course of development is quite different in the two cases. The tissue round the place where an animal has settled or where an egg has been fastened to the epidermis in this type begins to grow, rising up in the form of a fleshy mound or wall which continues to grow until the animal is wholly roofed in. The cavity in this case does not arise from an excava- tion (as in the embossed or “pocket” type), but from an overarching of the tissue. The 534 ALTERATION OF FORM BY GALL-PRODUCING INSECTS. external appearance of these galls is very varied. One of the simplest forms occurs on the leaves of the Ash (Fraxinus excelsior, see fig. 362°), where it is produced by the gall-gnat Diplosis botularia. The insect having laid its eggs in the chan- nelled depressions above the leaf-veins, fleshy cushions arise on either side of the groove which meet above and roof them over. The cushions of tissue forming the roof do not fuse; their succulent edges merely meet, and when the time comes for the gall-gnats to leave their temporary abode the tissue dries up and shrivels, 8 ( pally | D\\ Fig. 362.—Galls. ‘ Pine-apple gall on twigs of the Spruce Fir produced by the S i ieti: ; : pruce-gall Aphis (Chermes abietis). 2 Coverin aoe of the se a sae oe. byramidalis) produced by Pemphigus bursarius, 3 oa galls rene jal unus excelsior) produced by Diplosis botularia. 4 Covering gall on Pistacia (Pistacia Lenti: r a ; e tiscus) produced by Pem- phigus cornicularius. 5 Solid galls on the cortex of Duvaua longifoli rioaey I ngifolia produced by Cecidoses Eremii 6 itudi section of one these galls. 7 Capsule galls on the leaf of the Turkey Oak (Quercus Cerris) praduien segaaroee sce. leaving a gaping slit as shown in fig. 362°. The same thing happens on the leaves or rather leaf-veins of the Stinging-nettle (Urtica dioica) and of the Alder (Alnus glutinosa), where the galls are produced by gall-gnats (Cecidomyia urtice, alni) and on the midrib of Elm leaves (Ulmus campestris; see fig. 361°), where the alls are produced by a leaf-louse (Tetranewra alba). : The so-called turpentine gall-apples (Carobe di Giude; see fig. 3624), which SOLID GALLS. 535 are caused by leaf-lice on various species of Pistacia, also belong to the covering class. The rudiment of a foliage-leaf, which in the normal course of events would have developed into a pinnate leaf with dark-green elliptical leaflets, grows out into a pod-like structure not unlike a locust-bean (fruit of Ceratonia Siliqua). These galls are longitudinally grooved, and it can be seen more or less distinctly that the furrows correspond to the edges of the leaflets, only here the leaflets have become wrapped in, very much thickened and elongated, and fused with one another. In the cavity inclosed by the fused leaflets lives a colony of leaf-lice (Pemphigus cornicularius) which have developed under the protection of the gall. When it is time for them to leave the cavity the top of the pod opens by the separation and bending back of the tips of the fused leaflets which form the wall of the cavity (see fig. 362‘). The Chinese galls of commerce, produced also by Aphides (on Rhus semialata), develop much in the same way. They are hollow, irregularly pear-shaped structures with thin walls covered externally with a gray down. Two other covering galls which deserve special mention on account of their form arise on the petioles of the Poplar, particularly on the species Populus nigra, pyramidalis, and dilatata. The one, caused by a leaf-louse, Pemphigus bursarius (see fig. 362 *), consists of a smooth expansion, red in colour externally, on the upper side of the grooved petiole. If the local swelling be cut through it is seen to be hollow, the cavity in which the leaf-lice live being shut in by thick fleshy walls. The fleshy tissue of the walls is formed by a growth of the cells round the place where the gall-producing insect has settled. A hole is formed at a point remote from the petiole (where the growing tissue met and formed a dome) as soon as the time comes for the inhabitants to make their exit. This is bordered by thick lips as shown in fig. 362* The other gall which appears on Poplar petioles, pro- duced by Pemphigus spirotheca, is formed by the thickening of the edges of the grooved petiole, which rise up as fleshy cushions and meet above the depression. At the same time the petiole undergoes a spiral twisting, and a gall is thus pro- duced whose cavity is spirally twisted like the interior of a snail’s shell. The thickened edges of the petiole do not fuse; at first they fit close to one another, but later on they separate, and a spiral hole out of which the white, downy leaf- louse can creep is the result (see fig. 360’, p. 531). We will now leave the mantle-galls and pass on to a consideration of the solid or tubercular galls. These are of the nature of swellings of limited size on single plant-organs, and are produced by insects which pierce the plant-tissue and lay their eggs in the wound. In this way either the epidermis of the chosen spot alone is injured, or the egg is inserted into the deeper-lying tissues. In both cases an active cell-division is incited in the neighbourhood of the injury. If, however, the egg has only been deposited in the epidermis, the larva which arises from it must penetrate into the interior of the now swollen tissue; when the egg is laid at once deep down this farther penetration on the part of the larva is of course unnecessary. The cavity in which the larvee dwell may be called the larval chamber, and this sort of gall can be classified according to the number of chambers which it contains, 536 ALTERATION OF FORM BY GALL-PRODUCING INSECTS. whether only one or several (uf. figs. 363° and 363 7), A great variety is met with in the structure of the wall of the larval chamber. It always has a layer of juicy, thin-walled cells immediately surrounding the egg, known as the medulla or pith of the gall, and an outer layer which surrounds the inner like a skin or bark (see fig. 3607). In most instances a third layer is inserted between them which consists of very hard cells forming a protective layer. It should also be noted that the layers of the wall of the gall separate in many instances, so that it is possible to distinguish an “inner” and an “outer gall”. The gall-pith furnishes the larva with food when it emerges from the egg, and for this purpose the cells are stored with nourishing substances. The development of the pith takes place with great rapidity, and begins as soon as the egg has been laid in the tissue. The larva when hatched finds the inner wall of the chamber which has been fitted for its temporary abode always provided with the necessary food, and it immediately attacks and devours the juicy tissue with great avidity. The cells which are demolished, wonderful to relate, are replaced almost at once. The cells of the gall-pith remain capable of division as long as the larva in the chamber requires food, and the surface cells which have been devoured in the gall-chamber are soon replaced by new cells from below, just as grass which has been mown down or cropped by cattle in a meadow sends up new stems and leaves. The spheroidal gall arising on the leaves of Salia incana (ef. fig. 360°) has only one chamber, and here the larva lives at the expense of the starch and other food-materials contained in the extremely thin-walled cells which constitute the gall-pith (fig. 360"). The larva traverses the chamber in a circle, beginning the destruction of the cells at a certain place and eating on as it continues its peregrination (fig. 3601°). New cells have already been formed for its nourishment by the time it again reaches the place from which it started. The hard and cortical layers are modified in very many ways as protective measures against the drying up of the gall in summer on the one hand, and against the attacks of birds and larger animals on the other. For the latter purpose the cortical layer is often fashioned like the pericarps of fruits which have to protect the seeds (cf. p. 442). This also explains the bitter substances, hard skin, furry coat, bristling processes, and numerous other protective structures which are developed in and on galls just as on pericarps, and which contribute not a little to the remarkable similarity between galls and fruits. Many peculiar developments on the surface of these fruit-like galls cannot indeed be explained in this way, but, as in so many other cases, we conclude that they must afford some other advantage concerning which our understanding is still at fault. The external similarity between fruits and solid galls affords us useful points for classifying the latter into groups, which we may name berry-like, plum-like, apple-like, nut-like, capsule-like, &. The currant gall produced by Spathegaster baccarum on the male catkins of the Oak has not only the form and size of a Red Currant berry, but is also succulent and coloured red, and when several of these galls are formed on the same inflorescence it looks at first sight just as if racemes SOLID GALLS. 5387 of red currants had been borne by some chance or other on Oak twigs. The galls produced by the Beech-gall gnat (Hormomyia fagi) on the foliage of the Beech resemble small plums, being surrounded by a hard layer which consists of a stone kernel and a layer of cells which might be compared to the fleshy part of a plum. The galls caused by gall-wasps of the genus Aulaz on the nutlets of many Labiatz, especially on Nepeta Pannonica and Salvia officinalis also assume the form of stone-fruits. The insect lays its eggs in one of the four nutlets developed at the base of each flower; and within a week this grows into a smooth greenish- yellow ball which has the external appearance of an unripe cherry. A section through it shows that it possesses also the same structure as a cherry, plum, or stone-fruit in general. The succulent outer layer surrounds a hard stony kernel, but in the cavity of the kernel there lies the white larva of the gall-producer instead of the seed. These galls fall off just like fruits in July, and lie on the ground during the winter; and the mature insect does not bite an opening in the wall of the gall through which it can emerge until the following year. It has been already remarked at the beginning of this section how strong is the resemblance between apple-fruits and the spherical oak-galls, known as oak-apples, which are produced by various Cynipedes (see fig. 364°), together with the small red-cheeked galls produced by Rhodites Eglanterie and Nematus gallarwm (see figs. 361? and 3617) on Rose and Willow leaves respectively. Pith-galls which resemble certain dry fruits are very common. Those produced on the green cortex of young Oak twigs by Aphilothria Sieboldi (see fig. 3641) remind one of the fruits of species of Metrosideros, those produced by Neuroterus lanuginosus and Spathegaster tricolor on the leaves of the Turkey Oak (Quercus Cerris; see figs. 364" and 364714) have a decided similarity to the indehiscent fruits of the Wood- ruff and of the Goose-grass (Asperula odorata and Galium Aparine). The “spangle” galls produced on Oak-leaves by the gall-wasps Newroterus fumi- pennis and numismatis resemble the fruits of Omphalodes (see figs. 364" and 36418), and the galls on the leaves of Duvauwa longifolia produced by an insect Cecidoses Eremita have the form of a capsule which opens by an operculum (see figs. 362° and 362°). Like fruits these galls may appear in all imaginable con- ditions with smooth, warted, or rugged surfaces, or covered with woolly or velvety hairs, with bristles or spines, fringes or claws, or even with moss-like outgrowths. The galls with moss-like covering occurring on the Wild Rose have been known from remote times as Bedeguars. They are caused by the Rose-gall wasp (Rhodites Rose), which deposits its pointed, sometimes hooked eggs early in the spring in the substance of an undeveloped leaf while it is still folded up in the bud. The growth of the leaf becomes altered, the first sign being the development of numerous hairs. The larvee, when they creep out of the eggs, penetrate deeper into the tissue of the leaf, and it swells out into a solid gall contaming as many chambers as there are larve. Hairs and fringes continue to form on the exterior till those curious structures are formed which were said to have the power of inducing a peaceful sleep when laid under the pillow. Usually the stalks of the 538 ALTERATION OF FORM BY GALL-PRODUCING INSECTS. young bud-leaves are pierced and then the upper portion of the leaf becomes atrophied. More rarely is the egg laid in the epidermis of one of the leaflets, in which cage the leaves attain their normal size and only this particular leaflet is decorated with little bedeguars, as shown in fig. 3611. When the petioles of three young leaf-rudiments are pierced simultaneously, as often happens, three single galls are produced close together on a shortened axis, and the whole structure may then attain the size of a pine-cone. ; The portion of meristematic tissue which is pierced’ by the insect when it deposits its eggs sometimes remains an open passage; but more often a corky tissue is formed at the wounded spot which quite closes the chamber wherein the larva dwells. Under these circumstances the insect when it emerges must itself make an exit-passage from the gall, and this it does by biting a hole through it with its mandibles (see fig. 364°). The gall-wasps (Cynipedes) invariably leave the chamber which has hitherto served them both as a safe habitation and as an inexhaustible storehouse in this way. This does not occur, however, in some of those solid galls which owe their origin to gall-gnats of the genera Hormomyia, Diplosis, and Cecidomyia, for example, in those on the leaf-blade and petiole of the Aspen (Populus tremula) produced by Diplosis tremule and on the leaves of Willows (Salia Caprea, cinerea, grandifolia) by Hormomyia Capree. Here the exit-passage is formed during the development of the pith. The gall consists, as in most other solid galls, of a pith, a hard layer, and an epidermis, but the enormously developed pith and the hard layer do not quite entirely surround the small larval chamber, they leave a small aperture on the part of the gall which is most arched. As long as the epidermis stretches over this place the mouth of the passage is of course not evident, but when the time comes for the insect to quit the chamber a gaping slit is spontaneously formed in the tense epidermis. In many instances the insect or the pupa as it pushes forward may break through the thin skin. A peculiar closure which might be compared to a lid is formed in the common solid galls which are produced so abundantly on Beech leaves by Hormomyia fagi and which have been already alluded to. Just as the pupa of many Lepidoptera projects out of the hole in the cocoon which the caterpillar has spun for it far enough to allow the insect to fly away uninjured when it emerges, so that of Hormomyia fagi presses through the lid-like closure at the base of the gall, and the winged insect comes out leaving the chrysalis-case behind it. The opening of some solid galls, which resemble operculate capsules, and which may be termed capsule-galls, is especially remarkable and requires a more de- tailed description. As long as the larva or grub can remain and obtain food in the larval chamber the gall is completely closed, but when the time approaches for it to move its quarters and to enter the pupal stage in the ground a circular line of separation is formed in the tissue, and the part of the wall within the circle comes away as a lid. The process is seen very prettily in the gall produced on the leaves of the Turkey Oak (Quercus Cerris) by the gall-gnat Cecidomyia cerris (see fig. 362"). In its closed condition the gall is a firm rounded chamber DEHISCENCE OF CERTAIN GALLS. 539 so embedded in the leaf that it projects on the upper side as a small pointed cone, and on the lower side as a disc covered with a thick coating of hairs. In the autumn a circular piece like a lid becomes detached from the lower side of the chamber. It corresponds exactly with the extent of the hairy disc, and is so sharply defined that it looks as if it had been cut out with a knife (see figs. 362° and 362°). The operculum falls off, and the larva which had emerged from the egg and which has lived all the summer in the gall-chamber tumbles out and makes its way into the ground, where it begins to spin. By the next spring it has entered the pupal stage, and the gall-gnat creeps out of the chrysalis about May. Still more peculiar are the galls produced by Cecidoses Eremita on the green Fig. 363.—Solid Galls. 1 Capsule-like galls on a leaf of the Broad-leaved Lime (Tilia grandifolia) produced by Hormomyia Réaumuriana. 2 Longi- tudinal section through one of the galls, showing the maggot in the interior; x 2. 8% Longitudinal section through a capsule gall from which the inner gall is just being extruded; x 2. 4 Outer gall after the extrusion of the inner gall; x2. 5 Inner gall at the moment when the operculum is thrown off; x 2. § Capsule-galls on the leaf of a Brazilian species of Celastrus. 7 Longitudinal section through one of these galls; x 2. 8 The same after the inner gall has fallen out; x 2. 1 and 6 natural size. : cortical tissue of young twigs of Duvaua longifolia, a South American represen- tative of the Anacardiacee (see figs. 362° and 362°). The gall is quite spherical and very hard, and its large cavity conceals the caterpillar which has been hatched from the egg. When the time draws near for the formation of the pupa, a plug with a projecting rim is developed on the side of the gall furthest from its point of attachment. When the plug is pushed out a circular hole is left which leads into the gall-chamber through which the caterpillar escapes from its dwelling. People who have not seen these galls with their own eyes might almost think this description was the work of imagination. And yet there are still more wonderful forms in this class of gall-structures. On the foliage of the Lime (Tilia grandifolia) a growth arises round the eggs of the gall-gnat Hormomyia Réau- muriana which at first has the form of a flat lens inserted in the green tissue of the blade, but which gradually enlarges until it projects from the upper side like a 540 ALTERATION OF FORM BY GALL-PRODUCING INSECTS. blunt cone and from the lower as a hemispherical wart. The gall-chamber is inhabited by the maggot of the gall-gnat. The top of the conical part loses its colour in July and becomes yellow and brown, and a rim is formed around its summit. On cutting a vertical section through the gall at this stage it is seen that the tissue forming the wall of the chamber consists of two parts (see figs. 363”). The inner layer, which contains the maggot, is surrounded by an outer one which gradually passes into the green substance of the leaf and extends up to the rim just mentioned. The whole structure has separated into an “outer” and an “inner” gall, the inner gall resembling an egg lying in an egg-cup (¢/f. fig. 363”). During the summer the inner gall separates completely from the outer and is actually thrown off by it. For the accomplishment of this the tissue of the outer gall swells up very much, so that it exercises a pressure on the inner gall which is shaped not unlike a cone, somewhat narrower below than at the top (see fig. 363%). The extruded inner gall falls on the ground below the Lime-tree and assumes a dark- brown colour; the outer gall remains as a little crater embedded in the leaf-blade and ultimately shrivels up (cf. figs. 363! and 3634). The detached inner gall is smooth at the blunt and previously upper extremity, and striated at the other; it is not unlike a detached composite-fruit. The gall-gnat within feeds for a little time longer on the succulent lining, and then rests through the winter; in the spring it makes its escape. To do this it bites a ring-shaped groove below the conical top of the gall and presses against the roof, which, owing to the breaking of the tissues around the ring, comes away like a lid (see fig. 363°). A similar state of affairs prevails in a gall formed on the foliage of a Brazilian species of Celastrus (see figs. 363 ® 7 §), but here the inner gall (which comes away) has several chambers, and the outer gall has the form of a cup set in the green blade. The place of origin of all these solid galls depends of course upon the insects producing them. These are usually very fastidious about the place where they will lay their eggs, and it is truly astonishing with what care they search out spots difficult of access, and at once favourably situated as regards food supply and likely to afford a safe habitation for their offspring during the larval stages. The small gall-wasp Blastophaga grossorwm lays its eggs in the ovaries of the “ gall- flowers” in the interior of the figs of Ficus Carica (see p. 160 and figs. 240 * and 2407, p. 157). The gall-wasps Andricus amenti and Neuroterus Schlechtendali deposit them in the stamens of the Turkey Oak; the gall-wasp Cynips caput- medusce lays hers in the side of the bract-scales which surround the pistillate flowers of the Oak (Quercus sessiliflora and pubescens), and so produces a gall with innumerable stiff-pointed fringes entangled with one another which ward off the attacks of other animals (see fig. 364"). Countless gall-producing insects deposit their eggs on the lower side of foliage leaves, some preferring the lamina, others the veins. Andricus cwrvator prefers the margin of Oak leaves, Diplosis tremule the petiole of the Aspen at its junction with the blade. Several gall-wasps, as, for example, Andricus estivalis and Andricus grossularice, seek out the floral recep- tacle in the male catkins of the Turkey Oak for the deposition of their eggs, whilst ALTERATION OF FORM BY GALL-PRODUCING INSECTS. 541 several Cynipedes, e.g. Aphilotrix Sieboldi (see fig. 364) lay their eggs in the green cortex of the young twigs. Solid galls are very rare on roots, but they do occur Fig. 364.—Various Oak-galls. 1 Solid galls on the cortex produced by Aphilothria Siebolut. 2 Bud-gall from a foliage-bud produced by Cynips Hartigii. 8 Solid galls on an Oak twig produced by Cynips Kollari. 4 One of these galls cut in half. 5 Bud-galls from foliage-buds produced by Cynips lucida. §& One of these galls cut in half. 7 Leafy bud-galls produced by Aphilothria gemme. 8 Bud-galls from foliage-buds produced by Cynips polycera. 9% Longitudinal section through one of these galls. 10 Gall on the pericarp of Quercus pubescens produced by Cynips caput-meduse. 11-14 Spangle galls on a leaf of the Turkey Oak (Quercus Cerris); 11 produced by Neuroterus lanuginosus; 12 by Neuroterus numismatis; 18 by Neuroterus fumipennis ; 14 by Spathegaster tricolor. in this situation in the oak, being produced by the gall-wasps Aphilothria radicis and Biorhiza aptera. 542 ALTERATION OF FORM BY GALL-PRODUCING INSECTS. When several organs of a plant immediately adjacent to one another are con- cerned in the production of a gall it is said to be compound. Compound galls are for the most part produced from buds, and they are all comprehended under the general name of Bud-galls. They are extraordinarily varied in their characters, some being merely abbreviated axes clothed with scale-like leaves, in others only the base of the shoot is involved and above the gall it continues its growth quite normally, whilst in others again the axial portion of the structure is much swollen, and the leaves hardly represented at all. It is difficult to give any satisfactory classification of these bud-galls; still, for the sake of arranging our facts, we may distinguish these types, viz.:—the ordinary bud-gall, the cuckoo-gall, and cluster- gall. Ordinary bud-galls involve several, often all, the members of a shoot. The axis of the shoot is always deformed and abnormally thickened. The swollen portion contains in its interior one or several larval chambers surrounded by a pith-like layer. Two varieties of ordinary bud-gall may be distinguished. The first is leafless; no leaves are present, or, more correctly, they are transformed into tubercles, pegs, and knobs which merge insensibly into the swollen axis which contains the larval chamber. The second possesses leaves, the gall being covered with scale-like bracts or more or less fully developed green foliage-leaves. Amongst the leafless bud-galls the most interesting are those which are armed with special means of protection against the attacks of animals on the watch for the larve of the gall-wasps. The gall shown in figs. 364° and 364°, produced by Cynips polycera on the leaf-buds of Quercus pubescens and sessiliflora, which to a certain extent affects a whole lateral shoot, has the form of a young Medlar fruit, and on it may be seen 3-5 metamor- phosed leaf-structures projecting as stiff-pointed pegs which gradually pass into the tissue of the shoot axis. This gall is one-chambered, and the tissue of the wall has separated into an outer layer and an inner spherical pithy gall. The gall shown in fig. 364? is produced by the gall-wasp Cynips Hartigii which lays an egg in the middle of the leaf-bud of the Oak (Quercus sessilaflora). The bud does not develop into a leafy shoot, but into a small one-chambered gall with large tooth-like or club-like processes which represent metamorphosed leaves. The thickened angular ends of these projections fit closely to one another so as to form a sort of second outer coat to the gall-chamber through which hostile ichneumon-flies cannot penetrate. The gall much resembles the cone-fruit of a Cypress in the arrangement and form of its superficial processes. The galls produced from the buds of various Oaks (Quercus pendulina, sessiliflora, pubescens) by the gall-wasp Cynips lucida are still more peculiar (see figs. 364° and 864°). They contain several larval chambers with abundant pithy tissue, whilst innumerable slender processes resembling limed twigs in being very sticky on the capitate thickened end project from their exterior. Ichneumon-flies and other animals hostile to the gall-producers take good care not. to come into contact with these spikes which are to be regarded probably as trans- formed leaves springing from the swollen axis. Among the galls produced, from leaf-buds belonging to this group there are some in which the leaves are merely indicated as tubercles. This is the case, for example, in the many-chambered, BUD-GALLS. 543 spongy gall, red-cheeked on the sunny side but pale elsewhere, which is produced on the tips of the branches of the Oak by the gall-wasp Dryoterus terminalis, and looks very like a potato in shape. The leaves are only represented by small ill- defined knobs and ridges, just as in the potato. To this class of galls belongs also that to which the term “nut” is popularly applied, and even in commerce, the name has been transferred from this to the whole of the first group of compound galls (bud-galls). The “nut” is produced on the Oak by Cynips calicis as an angular and irregularly-grooved gall which originates at the end of a flower axis, and the cupule formed of several bract-scales as well as the ovaries are concerned in the growth. This class of galls also includes the irregular blunt swellings on Aspen twigs (Populus tremula), which are caused by the larva of a beetle (Saperda populnea), and in addition the many-chambered woody “canker cushions” as large as a nut which are produced on the branches of Willows by Nematus medullaris. The gall shown in fig. 3647, which arises on various Oaks (Quercus pedunculata, sessiliflora, pubescens) by the action of the gall-wasp, Aphilothria gemma, may be selected as a type of leafy bud-galls. It resembles the cone of a Hop or Larch, and is developed from a foliage-bud. It has a much-abbreviated swollen axis, whose tissue separates into an inner and outer gall, beset with numerous dry, brown lan- ceolate hairy scales having the form of bract-scales. Bud-galls which are covered with green foliage-leaves are produced by the gall-wasp Andricus inflator on the Oak, but they are more commonly met with on herbaceous plants, e.g. by Urophora cardui on Cirsiwm arvense, by Diastrophus Scabiose on several Knapweeds (Cen- taurea alpestris, C. Badensis, C. Scabiosa), by Aulax Hieracw on various Hawkweeds (Hieracium murorum, sylvaticum, tridentatum, &c.). Usually the foliage-leaves are stunted, and not infrequently the blades of some of them are quite obliterated, so that the gall in that region is only furnished with scaly leaf-sheaths. A Sage growing in the Isle of Crete so often bears leafy bud-galls resembling a small Quince-apple, produced by a species of Aulax, that Linnzus called it Salvia pomt- fera. The stem of this Sage is swollen out like a ball, and the spherical mass, covered with a gray felt of hairs on the exterior, is surmounted at the top with a group of small wrinkled leaves, which look like the persistent calyx of a Quince- apple. The best known and most widely distributed of these forms, found on the Hawkweeds named above, consist of knob-like swellings of the stem. The larval chamber is situated inside the enlarged pith, the ring of vascular bundles, which has undergone much shifting, forms the protective layer, and the cortex of the affected region of the stem forms the cortical layer of the gall. The epidermis is densely covered with hairs. Leaving the galls which consist of modified foliage-buds, we pass on to such as consist of metamorphosed flower-buds. They arise from flower-buds in which small gall-gnats have laid their eggs. The larva hatched from the egg lives in the cavity of the ovary, or.in one of its loculi when there are several, and this space, therefore, becomes the larval chamber. The corolla, which envelops the ovary in the flower- bud, remains closed, like a cap on the top of the larval chamber. The calyx becomes 544 ALTERATION OF FORM BY GALL-PRODUCING INSECTS. inflated, enlarged, and sometimes fleshy. The whole gall resembles a bud or small bulb; it is not unlike one of those bulbils which so often arise instead of flowers on the flowering axis of certain species of Allium. They occur especially on the Bird’s-foot Trefoil (Lotus corniculatus), where they are produced by the gall-gnat Cecidomyia Loti, on the various species of Mullein (Verbascum Austriacum, nigrum, Lychnitis, &e.) by Cecidomyia Verbasci, on several species of Germander (Teucrium montanum, Scordiwm, &ec.), caused by Lactomelopus Tewerri, and on the Rampion (Phytewma orbiculare), where they are produced by Cecidomyia phytewmatis. Closely allied to these bud-galls are those remarkable gall-structures which are commonly known in Austria as “cuckoo-buds”. The cuckoo is supposed to be concerned in their formation, just as it is in that of the frothy saliva-like masses deposited by the Cicada on the Cuckoo-flower (Cardamine pratensis). The name “euckoo-galls”” may be employed for the whole of this sub-group. They are char- acterized by their pale whitish colour, soft spongy tissue, and especially by the fact that they only involve the base of the shoot, while the upper end can continue its growth unaltered. In this respect they may be compared to a Pine-apple fruit, where the axis rises above the fleshy collective fruit (cf. p. 436) as a green leafy tuft, which does not lose its growing power even with the ripening of the fruit. The history of the development of cuckoo-galls is probably like that of covering galls; and the main distinction lies in the fact that in the former the gall is pro- duced not merely from a single organ or some part of it, but from a whole group of adjoining plant-members. The best known and most widely distributed gall of this group is produced by the pine-apple aphis Chermes abietis on the twigs of the Spruce Fir (Abies eacelsa, see fig. 3627, p. 534). Early in the spring, before the foliage-leaves have begun to unfold, the parthenogenetic females, the foundresses of the colony, attach themselves each to the base of a young leaf and lay a mass of eggs at the spot to which they have adhered. The larve, hatching, penetrate the surrounding parts of the shoot with their beaks; the shoot swells, as do the bases of the needles, and a growth, the Spruce gall or Pine-apple gall results. The gall somewhat resembles a small Fir-cone about an inch long, with the surface divided into small convex areas, each bearing a short needle-like projection in the middle; these are the deformed needles, which, becoming swollen, touch each other on the outside of the gall. They are separate inside, so that the gall contains a series of cavities or chambers. In these chambers the larve live in numbers, either entering the chambers during the growth of the gall or being inclosed by the swelling of the surrounding needles—this point is not certainly determined. They remain in the small cavities so formed and feed, cast their skins, and multiply there. In August the gall begins to dry up, each of the small cavities opens by a slit in front of the green needle-point surmounting the cushion (see fig. 3621, p. 584), and the winged insects now leave the place in which they have passed the spring and summer. Cuckoo-galls are met with almost as frequently on Stellate, viz. on various species of Bedstraw (Galiwm Austriacum, boreale, wliginosum, &¢.) and Woodruff BUD-GALLS. 545 (Asperula galioides, tinctoria, &.) as on Fir-trees. The infected parts of the shoot remain stunted, and white spongy cushion-shaped growths, which are somewhat grooved, arise at the bases of the leaves. Since the growing tissues of neighbouring leaves touch one another the grooves or channels form small cavities in which live the larves of the gall-producing gnats (Cecidomyia Galii and Asperulw). In the common Bedstraw (Galiwm Mollugo) these spongy growths arise, not from the bases of the leaves, but from the green cortex of the stem round the insertion of the leaves and lateral branches. They rise up as cushions and lobes, and several join together to form a sort of dome, under which the larve of the gall-gnat dwell. The foliage-leaves are scarcely altered in form, and when lateral twigs arise from the place they also are unchanged. It not infrequently happens that short lateral axes terminated by flowers spring up quite unmodified above the spongy white cuckoo-gall. Cuckoo-galls also occur on Crucifere, viz. on Barbarwa vulgaris, Nasturtvwm palustre, sylvestre, and Sisymbriwm Sophia. They are produced by Cecidomyia Sisymbrvi, and originate principally at the bases of the flower-stalks half-way up the inflorescences. They look like spongy white bodies which surround _ the pedicels like the brim of a hat. As the growths from neighbouring pedicels meet together they roof over chambers which serve as habitations for the larve of the gall-genats. Viewed from outside the galls appear like irregular white bodies inserted in the inflorescence, which remind one of the fruit of the white Mulberry- tree. The term cluster-gall is reserved for that type of bud-gall in which the axis is much restricted or stunted and covered with densely crowded leaf-structures; it is in the chinks and recesses between the crowded leaves of these galls that the insects concerned pass the whole or a portion of their lives. The animals which cause the galls belong to very different classes. Gnats, leaf-fleas, leaf-lice, and mites are the commonest varieties. The gnats only live in the galls during the egg and larval stages, but the others pass their whole life there. They invariably settle on the end of a shoot while it is still undeveloped in the bud. The axis of the shoot remains more or less stunted in consequence of the influence the animals exercise on it and its leaves undergo fundamental alterations. The blade or sheath of the leaf is deepened and hollowed to afford sufficient space to the animals which have established themselves between them, and as these parts of the leaves touch one another recesses are formed not unlike those which are developed in fir-cones for the growing seeds. The sheathing part of the leaf is often rather thickened, and its succulent cells serve as food for the animals living in the gall; in other instances the hollowed leaf-blades are thickly covered with hairs, and this coat then has the same significance with regard to the insects as the felt of hairs on isolated leaves already described. Very different forms of galls are produced according as to whether the free ends of the leaves turn back or remain in contact, and whether the axis from which the leaves spring is more or less contracted. Sometimes they remind one of open rosettes, sometimes of closed balls, bunches and tufts, sometimes of pig-tails and witches’ brooms. Vou. I. 85 546 ALTERATION OF FORM BY GALL-PRODUCING INSECTS. Clustered galls may be divided into two classes, those which develop in the region of the flowers and those in the foliage region respectively. The most noticeable and best known forms of the galls occurring in the foliage region on rudimentary leafy shoots are the following:—First, those peculiar structures on the tops of Willow twigs (Salia awrita, Caprea, grandifolia, &e.) which are popu- larly termed “Rose Willows”. They are caused by the gall-gnat Cecidomyia rosaria. The leaf-bud from which they arise keeps its axis quite short and develops on it numerous green leaves arranged like the petals of a double rose. The lowest leaves of the “rose” differ but slightly from the normal foliage of the particular species of Willow. Usually there seems to be only a shortening and broadening of the petiole and leaf-sheath, the green blade being almost un- altered. In the upper inner leaves, however, the sheath-like part of the leaf is much increased in size, and nearer the centre of the “rose” the leaves become scale-like, The leaf-blade has entirely disappeared, and the end of the contracted axis possesses only the remains of leaf-sheaths. It is worth noting that the number of leaves in a Rose Willow is always greater than would be found on an unaltered shoot of the same species. For example, if the number of leaves on the one-year-old shoot of the Sallow (Salia Caprea) is 25, the number in a “rose” on the same species would be at least twice as large. This can only be explained by supposing that a “prolepsis” has occurred, 2.¢. that not only the shoot laid down for the current year has developed, but also one originating from a bud of this shoot, which, under normal conditions, would not have developed until the following year. When autumn comes the rosette-shaped galls on the Willow bushes show up conspicuously at a distance because the leaves forming them do not fall off like the rest, but remain behind as brown dried structures at the ends of the branches. They are also found associated with the catkins. The rosette-shaped galls produced by the gall-gnat Cecidomyia crategi at the tips of Hawthorn twigs (Crategus Oxyacantha and monogyna) also claim atten- tion, They are full of bristles and resemble tiny birds’ nests. The stimulus of the gnat larvae excites a deeper and more frequent segmentation in the leaves and stipules. Narrower points and fringes which are much bent and which resemble the antlers of reindeers replace the broad lobes. Also soft spines with capitate ends rise up from the green cortex of the twigs and from the tissue of the leaf-blade, especially above the vascular bundles, and 3-5 of them often fuse together into cock’s-comb-like structures. These bristling rosettes on Hawthorn branches also remain long after the time the ordinary foliage falls off. In marked contrast to the rosette-like cluster-galls are others whose leaves all fold together in a ball something like the leaves of a cabbage, the whole gall having a button-like appearance. The outer leaves are round and hollowed on their upper side, and they usually fold together like mussel-shells, The inner leaves have a similar form, but they are much smaller and more concave, and they have become succulent and paler in colour. The galls produced by Cect- domyia genisticola on Genista tinctoria and those which Cecidomyia Veronice BUD-GALLS. 547 gives rise to on Veronica chamedrys, and which gall-mites produce on the Wild Thyme (Thymus Serpyllwm; see figs. 360+ and 360%, p. 531), form white buttons on the ends of the shoots which show up conspicuously from the dark green of the surrounding foliage. The white colour is due to the fact that the outer leaves, which fold together like mussel-shells, are thickly covered on the outside with white hairs. Cecidomyia Artemisie produces on the branches of Artemisia campestris a closed cluster-gall which is cased in white wool like a shroud. On the other hand, the large, button-shaped, closed cluster-galls which are produced by Cecidomyia rosaria on Willows (Salia purpurea, &c.) and by a gall-mite on the spikes of the Brome-grass (Bromus) are green and smooth, or at least they have not more than the usual number of hairs. On the shoots of the Yew (Taxus baccata), the Flax (Linum usitatissimum), Euphorbia Cyparissias, the Moss Campion (Silene acaulis), and several Ericas (Erica arborea, carnea, &c.) the influence of various gnats (Cecidomyia Taxi, Euphorbie, Erice, scoparice, &c.) produces galls with linear erect leaves crowded together into tufts. The base of the crowded leaves and the axis of the gall are usually rather thickened, so that it looks as if the linear leaves were set on a rounded button, and this is particularly marked in Huphorbia Cyparissias. This division includes the gall formations occurring on Juniper twigs (Juniperus communis), which are caused by the gall-gnat Lasioptera jwniperina. The acicular leaves of the Juniper are arranged in whorls of three on normal shoots. By reason of the influence of the gall-enat Hormomyia juniperina the whorls at the top of the twig become so changed that the last but one represents a cup bordered with three teeth in consequence of the broadening of the needles, while the terminal whorl is metamorphosed into a dwelling surrounded by three short leaflets. This gall closely resembles the cone of the Arbor Vite (Thuja occidentalis, orientalis, and plicata) in form. An insect, Livia Juncorum, produces galls on various Rushes (Juncus), espe- cially Juncus alpinus and lamprocarpus, which look like knots or tassels. The axis of the shoot is contracted, the sheathing portions of the leaves which cover one another are much widened, and the colour is pale except where it is reddened by exposure to the sun; their appearance is like the outer covering or top of a tassel. The stunted green blades which spring from the sheathing portions are thread-like and arranged as the loose strands of the tassel. Not infrequently short lateral shoots arise in the axils of some of the leaves, and then the whole structure looks like a bunch of tassels. Closely allied to these cluster-galls on the stems of Rushes are such as re- semble tufts and witches’ brooms, produced by mites on the branches of hairy Willows, especially on the white Willow (Salix alba). Instead of the long leafy Willow rod which would have emerged under ordinary circumstances from a foliage-bud, a confused mass of twigs with short leaf-scales is developed which at first seems a perfect mystery. By careful examination it is seen that the axis of the shoot laid down in. the bud has remained stunted, and that lateral 548 ALTERATION OF FORM BY GALL-PRODUCING INSECTS, shoots have developed from the axils of its leaves. These lateral shoots again develop lateral axes in the axils of their leaves, and so on to the third, fourth, and fifth degree. Thus, in the course of a month, shoots have unfolded, which, except for the influence of the gall-mites, would not have followed one another for three, four, five, or even six years, and therefore these galls afford us another instance of what has been termed “prolepsis” or precocious development of structures which would not yet arise. Of course all the axes of these shoots are dwarfed and the leaves which clothe them are diminished in size. The shortening and diminishing increase gradually, so that the axes and leaves of the fourth and fifth degree are much smaller than those of the second and third. ‘The last lateral shoots remain bud-like, and their small scaly leaves fold over one another like the bracts in the involucre of a Composite. The “ witches’ brooms” which are caused by gall-mites on Lilac (Syringa vulgaris) and Privet (Ligustrum vul- gare) bushes are similar in nature to these closed galls on the Willows. Frequently the metamorphosis of the leaves on the axes of the third, fourth, and fifth degree includes those’ of the floral region, and such cases form to some extent a bond of union between cluster-galls on foliage and on floral regions, respectively. One of the most remarkable changes exhibited by the gall-structures just men- tioned, viz. the abbreviation of the axis, is of course not to be noted in cluster- galls in the floral region. The part of the axis which forms the floral receptacle does not grow into an elongated shoot, but always remains short, and the floral- leaves it bears stand close to one another, forming whorls in whose niches and recesses numerous small animals can hide. But these animals effect other very marked alterations by their stimulus. In some flowers, instead of the norma’ red, blue, white, or yellow petals, green leaflets appear which resemble foliage- leaves in character, and then we say that the flowers have become “green” or “leafy”. In other plants the stamens are transformed into petals, and the flowers are said to be “double”. Finally, it may happen that the carpels which are usually united together to form a syncarpous ovary stand on the receptacle as distinct structures, and that to a certain extent their union has been dissolved. In these cases we speak of “antholysis” (ef. p. 80). The influence of gall-mites also produces metamorphosed flowers which may be both green and double, and in which the pistil may have separated into its individual carpels. The best flowers for observing these metamorphoses in all imaginable degrees are the small-flowered species of the Chickweed genus (Cerastiwm macrocarpwum, trwviale, &c.), several Caryophyllacese (Lychnis Viscaria, Saponaria officinalis, &e.), Crucifere (Cardamine uliginosa, Camelina sativa, Lepidiwm Draba), Gentians (Gentiana acaulis, rhetica), Speedwells (Veronica officinalis, saxatilis) and Milfoils (Achillea Millefolium, nana). In Speedwells the petals come to resemble leaves. The bunches, rosettes, and balls of small green leaves replacing the flowers are set close together on the rachis of the inflorescence and form green racemes and tufts, sometimes even small witches’ brooms. In Veronica sawatilis the rachis of the raceme, the pedicels, and the bracts are covered with hairs, which COMPOUND GALLS IN FLOWERS. 549 is not the case when the plants are free from the mites; the foliage-leaves in the neighbourhood of the raceme are also lobed and deeply indented, which again is not the case in uninfected plants of this species. In the capitula of the above- named Milfoils the peripheral ray-florets as well as the central tubular ones become leaf-like, and this gives rise to the most peculiar forms.