Part 1
YeWl- Joe ; ANNALS OF BOTANY VOL. XX Orford PRINTED AT THE CLARENDON PRESS BY HORACE HART, M.A. PRINTER TO THE UNIVERSITY ANNALS OF BOTANY EDITED BY ISAAC BAYLEY BALFOUR, M.A. M.D, F.R.S. KING'S BOTANIST IN SCOTLAND, PROFESSOR OF BOTANY IN THE UNIVERSITY AND KEEPER OF THE ROYAL BOTANIC GARDEN, EDINBURGH DUKINFIELD HENRY SCOTT, M.A., LL.D., F.R.S. LATELY HONORARY KEEPER OF THE JODRELL LABORATORY, ROYAL BOTANIC GARDENS, KEW JOHN BRETLAND FARMER, D.Sc, M.A., F.R.S. PROFESSOR OF BOTANY, ROYAL COLLEGE OF SCIENCE, LONDON AND WILLIAM GILSON FARLOW, M.D. PROFESSOR OF CRYPTOGAMIC BOTANY IN HARVARD UNIVERSITY, CAMBRIDGE, MASS., U.S.A. eww ASSISTED BY OTHER BOTANISTS a ar vw wo %%y VOLUME XX sole. With Thirty-five Plates, Map, and Forty-three Figures in the Text esveusd Bondon HENRY FROWDE, M.A., AMEN CORNER, E.C. “~~ OXFORD: CLARENDON PRESS DEPOSITORY, 116 HIGH STREET 1906 a a ed wee & & howe Te » * & - - wwe © & es - ww . . ww * ee ww » ov ew » * we wy, Ld - ". w wv - * e®eave wsesetbe . » ~w 7 @ ooe ew - ~ . . oe” wuvese e . 103638 CONTENTS. No. LXXVII, January, 1906. PLowMAN, A. B.—The Comparative Anatomy and Phylogeny of the Cyperaceae. With Plates I and II and two Figures in the Text. BLACKMAN, V. H., AND FRASER, Miss H. C. I.—Further Studies on ‘the Sexuality of the Uredinese. With Plates III and 1V . : . . . . . . Buiuer, A. H. R.—The Enzymes of Polyporus squamosus, Huds. . . Ponb, R. H.—The Incapacity of the Date Endosperm for Self-digestion Ewart, A. J.—The Influence of Correlation upon the Size of Leaves. With two Fi igures in the Text . . HuMmpurey, H. B.—The Development of Fossombronia longiseta, “Aust. With Plates v and VI and eight Figures in the Text . . . . . . NOTE. Scott, D. H., AND MASLEN, A. J.—On the Structure of Trigonocarpon olivaeforme No. LXXVII, April, 1966. VINES, S. H.—The Proteases of Plants. IV. . . . . THISELTON-DyER, Sir W. T.—Morphological Notes, With Plates VII- IX : WorsDELL, W. C.—The Structure and Origin of the Cycadaceae. With seventeen Figures in the Text . Hii, T. G.—On the Seedling Structure of certain Piperales. “With Plate x and ‘three Diagrams in the Text Breer, RUDOLF.—On the Development of the Spores of Helminthostachy reylanica. With Plates Xl and XII . . SALMON, E. §S.—On Ojidiopsis taurica (Lév. ), an “ endophytic member ‘of the Exysiphaceae. With Plates XIII and XIV. Ewart, A. J., AND MASON-JONES, A. J. The Formation of Red Wood i in Conifers. ‘With Plate XV . . STaPF, Orro.—The Statices of the Canaries of the Subsection Nobiles. I NOTE. RIDLEY, H. N.—On the Foliar Organs of Monophyllaea No. LXXIX, July, 1906. ARBER, E. A. NEWELL.—On the Past History of the Ferns. With a Diagram in the Text Ex.is, DAVID.—The Life-history of Bacillus hirtus (Synonyms Bacterium hirtum, Henrici ; Pseudomonas hirtum, Ellis). With Plate XVI ROBERTSON, AGNES.—Some Points in the Morphology of Phyllocladus alpinus, Hook. “With Plates XVII and XVIII . . HILL, T. G.—On the Presence of a Parichnos in Recent Plants. W ith Plates XIX and XX . Breer, RuDOLF.—On the Development of the Spores of Riccia glance. With Plates XXI and XXII Boop.e, L. A.— The Monoecism of Funaria hygrometri, Sith. Ww ith four Figures j in the PAGE 35 49 61 79 83 109 113 123 129 161 177 187 201 205 313 215 233 259 267 275 393 Vi Contents. PAGE STAPF, OTTO.— The Statices of the Canaries of the Subsection Nobiles. IJ. With a Map in the Text . . . . . 30! Yarp, R. H. —Frnit-dispersal in Adenostemma viscosum. A Biological Study. With Plate XX. . . . . . . . . . . . . . . gil NOTES. Scott, D. H.—The Strocture of Lepidodendron obovatum, Stemb. . . . . . 317 Boopie, L. A.—Lignification of Phloem in Helianthus . . . . . , , - 319 CAMPBELL, D. H.—Germination of the Spores of Ophioglossum . ; ; . . . 3a! Multiple Chromatophores in Anthoceros . , . ; ; . - 32a No. LXXX, October, 1906. Prain, D.—A Review of the genera Meconopsis and Cathcartia. With PlatesXX1]VandXXV_ 323 Sewarp, A. C.—Anatomy of Lepidodendron aculeatum, Stemb. With Plate XXVI and three Figures in the Text . . . . , . . 371 Jutergy, Epwarp C.—The Wound Reactions of ‘Brachyphy!lum. With Plates XX VII and XXVHI. . 383 Hitt, AktTHUR W.—The Morphology and Seedling Structure of the Geophilous Species of Peperomia, together with some Views on the Ongin of Monocotyledons. With Plates XXIX and XXX and three Diagrams inthe Text. «www. 898 ROBERTSON, .\GNES.—The ‘ Droppers” of Tulipa and Erythronium. With Plates XXXI and XXXH_. . ; . . . . . . . . ; . 429 Hotm, THZODORE.—Bartonia, Muehl, An Anatomical Study. With Plates XXXIII and XXXIV . , . . . . . . ; . . 441 Prince, Georce J.—Studies of Irritability in 1 Plants. With Plate XXXV_. . . - 449 NOTES, Hemsiey, W. Bottinc.—On the Julianiaceae, a new Natural Order of Plants. (Abstract.) . 467 Hitz, T. G., AND Dg Fraine, E.—On the Seedling Structure of Gymnosperms . - 47! Hitt, T. G.—On the Seedling Structure of certain entrospermac . , . ; . 473 CONTENTS. . , . , . . , . v INDEX. ; ; ; . , . . , . . . . vii aos INDEX. A. ORIGINAL PAPERS AND NOTES. ArBeEr, E. A. NEWELL.—On the Past History of the Ferns. With a Diagram in the Text Breer, RuDoLF.—On the Development of the Spores of Helminthostachys zeylanica. With Plates XI and XII . . ————— —— On the Development of the Spores of Riccia glauca. With Plates XXI and XXII . . BLACKMAN, V. H., AND FRASER, Miss H, C. 1.—Further Studies on the Sexuality of the Uredinese. With Plates I and IV . BooDLe, L. A.—The Monoecism of Funaria hygrometrica, Sibth. With four Figures i in the Text. . . . ———______—— Lignification of Phloem i in Helianthus . Bu iver, A. H. R.—The Enzymes of Polyporus squamosus, Huds. CAMPBELL, D. H.—Germination of the Spores of Ophioglossum ---- - - Multiple Chromatophores in Anthoceros de FRAINE, E., see Hill, T. G. Ex.is, DAVID. —The Life-history of Bacillus hirtus (Synonyms Bacterium hirtum, Henrici ; Pseudomonas hirtum, Ellis). With Plate XVI . . . . . Ewart, A. J.—The Influence of Correlation upon the Size of Leaves. With two Figures in the Text . Ewart, A. J., AND MASON-JoNEs, A. J—The F ormation of Red Wood i in Conifers. With Plate XV. . . . . . . . . . . Fraser, Miss H.C. I., see Blackman, v. H. HemMsvey, W. BOTTING. —On the Julianiaceae, a new Natural Order of Plants. (Abstract.) . HILt_, ARTHUR W.—The Morphology and Seedling Structure of the Geophilous Species of Peperomia, together with some Views on the Origin of Monocotyledons. With Plates XXIX and XXX and three Diagrams in the Text . : Hitt, T. G.—On the Seedling Structure of certain Piperales. With Plate x and three Diagrams in the Text . On the Presence of a Parichnos i in Recent Plants. With Plates XIX and XX. - —— — On the Seedling Structure of certain Centrospermae . . Hitt, T. G., AND DE FRAINE, E.—On the Seedling Structure of Gymnosperms . Hoi, THEODORE.—Bartonia, Muehl. An Anatomical Study. With Plates XXXIII and XXXIV . . . Humpnrey, H. B.—The Development ‘of F ossombronia longiseta, Aust. “With Plates v and VI and eight Figures in the Text . Jerrrey, EpwarpD C.—The Wound Reactions of Brachyphy/m. With Plates XXVII and XXVIII. . . . , . . Masten, A. J., see Scott, D. HL. Mason-JOnes, A. J., see Ewart, A. J. Peirce, GeorGE J.—Studies of Irritability in Plants. With Plate XXXV . . , Prowman, A. B—The Comparative Anatomy and Phylogeny of the CyPeracene: With Plates I and II and two Figures in the Text . . . . Ponb, R. H.—The Incapacity of the Date Endosperm for Self-digestion , . . , PRAIN, D.—A Review of the genera Meconopsis and Cathcartia. With Plates XXIV and XXV . . , . . . . . Rrp._ey, H. N.—On the Foliar Organs of Monophyllaea . ROBERTSON, AGNES.—Some Points in the Morphology of Phyllocladus alpinus, Hook. With Plates XVII and XVIII . . -—— —— -—— —-- The‘ Droppers of Talis and Erythronium. Ww ith Plates XXXI and XXXII. . . . . . . ‘ : PAGE 215 177 375 35 293 319 49 321 324 233 79 201 467 395 161 267 473 471 83 383 449 61 323 213 259 429 eee vill Index. PAGE SALMON, E. S.—On Oidiopsis taurica (Lév.), an endophytic member of the Reyiphaceas. With Plates XIII and XIV. . 187 Scott, D. H.—The Structure of Lepidodendron obovatum, Stemb. . 317 Scott, D. H., AND MASLEN, A. J.—On the Structure of Trigonocarpon olivaeforme 109g SEWARD, A. C -—Anatomy of Lepidodendron aculeatum, Sternb. With Plate XXVI and ‘three Figures in the Text . 371 STAPF, OTTO.—The Statices of the Canaries of the ‘Subsection Nobiles. I. . . . 205 The Statices of the Canaries of the Subsection Nobiles. II. With a Map in the Text . . : . . 3or THISELTON-Dygr, SIR W. T. —Morphological Notes. ‘With Plates VII-IX 123 VINES, S. H.—The Proteases of Plants. IV . 113 WoRSDELL, W. C.—The Structure and Origin of the Cyeadacone “With seventeen Fi igures in the Text... 129 Yaprp, R. H. —Fruit-dispersal in " Adenostemma. viscosum. A ‘Biological Study. ‘With Plate XXIII. : . . : . . . . grr B. LIST OF ILLUSTRATIONS. @. PLATES. I, 1I. Anatomy of Cyperaceae (PLOWMAN). Ill, IV. Uredineae (BLACKMAN and F RASER). V, VI. Fossombronia (HUMPHREY), VIL-IX. Protective Adaptations (THISELTON-Dykr). X. Piperales (T. G. HILL). XI, XII. Helminthostachys (BEER). XIII, XIV. Oidiopsis taurica (SALMON). XV. Red Wood in Conifers (EWART and MASON-JONES),. XVI. Bacillus hirtus (ELLIS). XVII, XVIII. Phyllocladus alpinus (ROBERTSON). XIX, XX. Parichnos (T. G. H1Lt). XXI, XXII. Riccia glauca (BEER). XXIII. Adenostemma viscosum (YAPP). XXIV, XXV. Meconopsis and Cathcartia (PRAIN). XXVI. Lepidodendron aculeatum (SEWARD). XXVII, XXVIII. Wound Reactions of Brachyphyllum (JEFFREY). XXIX, XXX. Peperomia (A. W. HILL). XXXI, XXXII. ‘Droppers’ of Tulipa and Erythronium (ROBERTSON). XXXII, XXXIV. Bartonia (HOLM). XXXV._ Irritability in Plants (PEIRCE). 6. WoopcutTs. 1, 2. Comparative Anatomy and Phylogeny of Cyperaceae (PLOWMAN) . 32, 33 I, 2. Influence of Correlation upon the Size of Leaves (EWART) . - 80 1-8. Development of Fossombronia longiseta (HUMPHREY) . 84-6, 89, 90, 93, 100, Tol 1-17. Structure and Origin of Cycadaceae (WORSDELL) . 136-42, 144, 145, 147, 149-52 I-IJI. Seedling Structure of certain Piperales (T. G. Hitt) 162, 164, 168 1. Past History of the Ferns (ARBER) . 229 1-4. Monoecism of Funaria hygrometrica (Boobie) 295, 296 Map. Statices of the Canaries of the Subsection Nobiles (STAPF) 309 1-3. Anatomy of Lepidodendron aculeatum (SEWARD) . . . 374-6 1-3. Morphology and Seedling Structare of Geophilons Species of Peperomia (A. W. HILL). 402, 418, 420 4 The Comparative Anatomy and Phylogeny of the Cyperaceae'. | BY AMON B. PLOWMAN. With Plates I and II and two Figures. SYNOPSIS. PAGE 1, INTRODUCTION. 2 2. PROBLEMS F 2 3. THE Root ° 3 4- THE RHIZOME . . . . . : . . . . . : 4 a. General Morphology . . . . . . . . - oe 5 46. The Amphivasae . . . 6 c. The Centrivasae . . 7 d. Generic Key . . . . ° 9 e. Descriptive Table . . . 12 5. THE AERIAL STEM . : . . . . . . . . . . - «10 a. General Morphology . - © 6 II 6. Mode of Origin and Apical Growth . II c. Internal Anatomy . II aa. The Chlorocy peraceae 14 66. The Sclerocyperaceae . 15 ce. Occurrence of Cambium 18 @. Descriptive Table . . . . oe 16 e. Nodal Complexes. . » 18 jf. Course of Fibro-vascular Bundles 18 g- Cortical Bundles. . 22 hk. The Scapose Type . 21 8. Axillary Buds 2 6. THe Lear... . . , . . 2 . . . . - 23t a. General Morphology . . . . - . . - «6 2 6. Internal Anatomy . 22 ¢. Mode of Insertion . . 22 d. Dominance in Stelar Development . 23 7. THE FLORAL AXIS ; . 23 a. Morphology and Anatomy ° 23 4, Dicotyledonous Arrangement of Bundles . 23 8. Fossil CYPERACEAE . 24 g. PHYLOGENETIC CONSIDERATIONS . | a. Inter-relationships of the Cyperaceae . . . . . . oe - e 35 6. Position among the Monocotyledons . 26 ¢. Bearing upon the Phylogeny of the Monocotyledons 26 Jo. SUMMARY AND CONCLUSIONS. 27 18. LITERATURE . - 28 12. EXPLANATION OF THE PLATES 30 1 Contributions from the Phanerogamic Laboratories of Harvard University, Ne. 2. (Annals cf Botany, Vol. XX. No. LEXVII. January, 1906.) B 2 _ Plowman.—The Comparative Anatomy and INTRODUCTION. INCE the publication of De Bary’s (12) great work on Comparative Anatomy, and Van Tieghem’s (57) subsequent ‘stelar theories,’ there has accumulated a considerable body of literature devoted to this phase of botanical study. The work of De Bary marked the culmination of that earlier mode of treatment which considered the axis of vascular plants to be made up of more or less numerous, wholly independent fibro-vascular bundles, embedded in a common matrix of fundamental tissue. Van Tieghem (57) later introduced the idea of a fundamental stelar unity of the fibro-vascular system, and showed that this unity held essentially true of all the Vasculares. The elaboration of this view lent new interest and broader possibilities to the anatomical study of plants; and while many of the details and assumptions of Van Tieghem’s theory have been disproved or modified by the later investigations of Gwynne- Vaughan (20), Jeffrey (27), Boodle (4), Faull (18), Worsdell (61), Schoute (45), Chrysler (9), and others (5, 17, 21), yet the important fact still remains that in the fibro-vascular system of -higher plants we find an essential unity of plan which is fairly comparable to that found in the skeletal system of vertebrate animals, and which is apparently of hardly less importance from the phylogenetic point of view. . In the hope of gaining some new light upon the vexed question of the relationships of the Monocotyledons, the writer has undertaken the present study of the Cyperaceae, a group which is especially interesting on account of its apparently somewhat isolated position and its clearly-marked characters. Upon making a general survey of the problem before us, the following questions, among others, present themselves for careful consideration, if not, indeed, for solution :— 1. Anatomically considered, do the Cyperaceae constitute a clearly- defined group? 2. What subdivisions of the Cyperaceae may be made upon the basis of anatomical characters ? 3. Of what importance are oecological factors in determining the anatomical features of the group? 4. What is the nature, origin, distribution, and significance of the amphivasal bundles found in this group? 5. What is the course of the fibro-vascular bundles through the stem ? 6. Of what importance are leaves as factors in stelar development? 7. What structural characters of the Cyperaceae are ancestral, and what are more recently acquired ? 8, What are the probable phylogenetic inter-relationships of the Cyperaceae? Phylogeny of the Cyperaceae. 3 g. What is the position of the Cyperaceae among the Monocoty- ledons? 10, What bearing have the observed facts upon the phylogeny of the .Monocotyledons as a whole? | Before entering upon a discussion of these problems it will be neces- Sary to consider in more or less detail some of the structural characteristics of representative genera and species of the Order. THE ROOT. The internal structure of the root in the Cyperaceae presents a ‘striking degree of uniformity. The central cylinder is compact and rela- tively small, surrounded by a well-marked endodermal sheath, the cells of which commonly have the inner wall much thickened. The medulla is ‘composed of round, more or less sclerotic cells of relatively small size, with -sharply-defined intercellular spaces. The xylem masses are radially -disposed within the bundle-sheath, alternating with the phloem masses. Their number is variable, but rather smaller than that found in the majority ‘of Monocotyledons, ranging from six to eight in species of Eleocharis to fifteen to eighteen in certain Carices, while, according to Kny (32), these numbers may approach or even exceed 100 in the roots of some Palmales. ‘Each xylem-mass consists of a few short, small, thin-walled tracheids -arranged in a radial plate terminated centrally in a single relatively large vessel. It frequently happens that there are not as many large vessels as there are xylem-masses, in which case growth pressure may cause a vessel to lie at the point of a V of which the arms are two distinct xylem-masses, ‘which are brought in this way to subtend the intervening phloem-mass, in ‘a manner remotely suggestive of the collateral arrangement of the elements ‘found in the stem. It is worthy of note that,.as shown by Klinge (31), the elements of the protoxylem are as a rule neither spiral nor annular, but scalariform, reticulate or pitted. . The phloem is less conspicuously developed, the masses being quite ‘small and often but slightly differentiated. .In mature roots the elements are either partially lignified or considerably crushed by growth pressure. It has been shown by Chauvaud (8) that the sieve-tubes are derived in each case from a ‘ pericambial’ cell lying next to the endodermis. The phloem is separated from the xylem by a few or sometimes numerous rows of “usually somewhat sclerotic parenchyma-cells. The cortex of the young root consists of many layers of thin-walled cells arranged with great regularity in radial rows. In mature roots