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SYSTEMATIC ANATOMY OF THE DICOTYLEDONS

A HANDBOOK

FOR LABORATORIES OF PURE AND APPLIED BOTANY
BY

DR. HANS SOLE REDER
PROFESSOR OF BOTANY IN THE UNIVERSITY OF ERLANGEN

TRANSLATED
ltOVAL BOTANIC

BY

L. A. BOODLE, F.L.S.
GAil DENS, KEW

AND

F. E. FRITSCH, D.Se., PH.D., F.L.S.
ASSISTANT PltOFESSOIt OF BOTANY, UNIVItItSITY OF LONDON, UNIVEIlSITY COLLEGE AND LECTUUIt IN BOTANV, L\ST LOIfDON COLLEGE, UNIVEI.SITY OF LONDON

REVISED BY

D. H. SCOTT, M.A., LL.D., PH.D., F.R.S.
LATE HOIfOI.AltV ItItItPEI. OF THE JODULL KEW L4BOllATOl.V I.OYAL BOTANIC GARDENS,

VOL. I
INTRODUCTION, POLYPETALAE,GAMOPETALAE
WITH US FIGURES IN THE TEXT

OXFORD AT THE CLARENDON PRESS
1908

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A.HENRY FROWDE. PUBLISHER TO THE UNIVERSITY OF OXFORD LONDON. EDINBURGH NEW YORK AND TORONTO Digitized by Google . M.

L. DR. RADLKOFER SEINEM HOCHVEREHRTEN LEHRER UND VORSTANDE 'tHIS BOOK IS DEDICATED BY THE AUTHOR Digitized by Google .TO PROF.

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each of whom has hitherto often remained too much isolated in his own department. by some of our greatest systematists. with marvellous completeness and exactitude. the study of anatomical structure in relation to taxonomy has been but little pursued in this country. though a beginning was made. much facilitate the introduction of modern anatomical methods into the daily work of the Herbarium and the Economic Museum. both the Addenda and the Concluding Remarks having received from the author's hand all the emendations which will be embodied in the new German edition. The work is universally recognized among Botanists as the standard authority on the subject of the anatomy of plants as applied to systematic purposes. The case for the anatomical method is put by the author with singular judgement and moderation. as will be seen from the Introduction and the Concluding Remarks. whether their bent be systematic or otherwise. they may be read with interest and advantage by all students of botany. Until recently. explain so clearly the object and scope of his book that it is unnecessary to enter on any further detailed exposition of its purpose. The latter form a very full and valuable summary of the various anatomical characters and their value in classification. Digitized by Google . it is hoped. The present translation will. many years ago. Professor Solereder's book must serve as the type of all treatises dealing with the structure of other classes from a similar point of view. to the original German edition and to p the present English translation. Covering.EDITORIAL PREFACE PROFESSOR SOLEREDER'Srefaces. the ' Systematic Anatomy' forms an indispensable work of reference to all engaged in systematic studies. as it does. It may be pointed out that the English edition has been brought thoroughly up to date. though limited to the Dicotyledons. the most extensive and important division of the Vegetable Kingdom. and serves at the same time as by far the best source from which students may gain an idea of the objects and methods of this young and vigorous branch of Botany. One of the advantages of the study of systematic anatomy is that it brings together the work of the laboratory botanist and the taxonomist.

which came in too late to be appended to the lists under each Natural Order in the Addenda. Fritsch. first begun by Mr. are to be congratulated on the successful completion of their labours. Some additional references to the literature. with their German originals. Fritsch alone. the work of translating the Addenda and Concluding Remarks fell to Dr. and the translators. and in the necessary consultations with the translators on all questions that arose in the course of their work. embracing such a vast amount of minute and accurate detail. H. 1908. at the end of the second volume. will be found collected at the end of the book. I have aimed at securing a sufficient uniformity of style throughout the book. and have satisfied myself that the translation faithfully conveys the author's meaning. SCOTT.vi PREFACE The work of translating so extensive a treatise. My own part has consisted in the revision of the proofs throughout. Digitized by Google . The rendering of technical terms has in some cases presented a certain amount of difficulty. The translation. Boodle. was carried out jointly by him and his colleague up to the end of the. Boodle and Dr. it is proposed to add a glossary of the less familiar terms. has naturally been extremely arduous . Mr. Marc" 31. D. Natural Orders.

SOLEREDER.' which give a survey of the anatomical characters of the vegetative organs and of the distribution of these characters among the Dicotyledons. which have not yet been published elsewhere. under the respective Orders. 1-903 of the German edition). 1901. compiled at the request of my English colleagues. EIlLANGIlN.-to all the colleagues who have assisted me in my work by the forwarding of separate copies or of material for investigation. Digitized by Google . The Orders are dealt with in the sequence of Bentham and Hooker's system. who are so necessary for its further elaboration and obligatory introduction into all systematic researches (monographs. questions of affinity. is followed by an appendix. introduced from time to time.' originally published in the year IB99 by F. contains the translation of the Introduction and of the Anatomical Characters of the Dicotyledonous Orders (pp. 904-79).PREFACE BY THE AUTHOR EDITION TO THE ENGLISH THE present English edition of the 'Systematische Anatomie der Dicotyledonen.-last not least. exactly the same plan has been followed in the Addenda as in the principal part of the book (see the preface to the German edition). and. As regards the arrangement of the subject-matter and the terminology of the species. which summarizes. the results of recent researches. This. Readers who are conversant with the subject will also find new observations. At the same time I wish to tender my heartiest thanks to the translators for the great labour expended on the preparation of this work. establishment of new genera and species). In the sections devoted to the structure of the leaf and the structure of the axis the anatomical characters are discussed in the same serial order as in the principal part of the book. for it carries with it the hope that the anatomical method will find new adherents. In concluding. I should like above all to express the pleasure which the translation of my book has afforded me. In the descriptions of the individual Orders supplementary details to the' review of the anatomical features. and constitute a revised treatment of the' Schlussbemerkungen ' of the original German edition (pp. of which I was acquainted with the title only or of which only abstracts were available. are again placed in square brackets. papers.' the' structure of the· axis' and in some cases also the' structure of the root' succeed one another.' the' structure of the leaf. H. Enke in Stuttgart. as well as to the Delegates of the Clarendon Press for their invariable readiness to meet my wishes. A considerable amount of trouble has been taken in completing the references to literature. Oc/~. the principal part of the book. The book terminates with the • Concluding Remarks.

Polypompholyx Biovularia • 589 591 591 592 583 601 613 624 627 628 630 599 597 597 598 594 6II Zombiana 642 636 1. CamJ?anulaceae (incl. . . Digitized by Google . • Columelliaceae Gesneraceae Bignoniaceae Pedalineae Acanthaceae Myoporineae Selagineae . • . Comaceae . .xii POLYPETALAE CONTENTS (continued) PAGE OF VOLUME I (continued):PAGE ANATOMICAL CHARACTERS OF THE DICOTYLEDONOUS ORDERS Cucurbitaccae Begoniaceae Datisceae .. Compositae . Verbenaceae Labiatae • _ P1antagineae Pinguicula . . . Calycereae . • Ericaceae . CandoUeaceae (Stylidiaceae) Goodeniaceae. . Genlisea . . . Cacteae . . 528 534 538 547 548 552 554 562 573 573 575 581 526 550 Lonchostoma . .. Utricularia. Lobeliaceae) Vacciniaceae . . Valerianeae Dipsaceae . . . GAMOPETALAE 406 415 419 426 432 406 389 398 Salvadoraceae Apocynaceae Asclepiadeae Loganiaceae Caprifoliaceae Rubiaceae . Cuscuteae Nolaneae Solanaceae Retna and Scrophularineae Orobanchaceae Lentibularieae Des/ontainea and Plocosperma . 439 444 454 455 456 456 469 47 I 473 476 479 489 490 494 494 495 501 507 512 516 519 521 Gentianeae Polemoniaceae Hydrophyllaceae Boragineae Convolvulaceae. Umbelliferae Araliaceae . Ficoideae . Monotropeae Epacrideae Diapensiaceae Lennoaceae P1umbagineae Primulaceae Myrsineae. . . Sapotaceae Ebenaceae Styraceae Oleaceae . .

in NatiirL PftlDmlgm.-H. in OQomeles bordered pitting is. pp. Reslnoc:ylteD. Tell. 2. Bot. intemaz. de BuiteD&org. t. Perldriileu. 1 For Od#mlks see also Warhurg. xviii. MObilIS. which are composed of a multiseriate stalk of variable length and ora spherical or ellipsoidal. The Cacteae. 382). se. :nii. DilL. glandular head. AnaL d. Schoeoett'. Of this small Order.. Soc. CeutralbL 18g6.-Doclwtre. I3--Petit. AmI. only Datisca cannabina. at least in some cases.406 BEGONIACEAE Lem4i_. Abt. The cork of Octomeles consists of cells with wide lumina and relatively thin walls. Bot. transverse walls with simple perforations. et nat . Akad. 1887. 125-7 and Fig. for in the other genera a fleshy shoot. d. pp.-Wittlin. In Octomeles sumatrana groups of stone-celJs. Tell. 1892p 239 pp.-Warbarg. deutsch. Topogr. while the adjacent wall of the medullary ray is thickened in a well-marked reticulate manner. poln. Kopemilt in Lemberg. Regarding the structure of the leaf. which are found in Octomeles. phys. 1887. pp. Abt. 11194. this shoot takes over the function of the foliage leaves. 6 .. 46. voL mi. 50-5 (discaaiOll of Schoeoett's tIeatise in Ko. iii.pp. 18940 p. MiiDcheD. The vascular bundles oi the veins are accompanied by mechanical tissue in Octomeles sumatrana only.. Paris. constitute an Order which is distinguished from all others by its habit. 151. in Nltilrl.. multicellular. Bar Ie IIOIIB-geIIIe E. Secondary groups of bastfibres are present in Octomeles.mOl. the mesophyll of this plant contains spicular cells. which includes three genera with five species. circular shield with an entire margin. Ber. The trichomes occurringin Datisca are glandular shaggy hairs. Nebeuz. po loo. Foum. u. 18g7. 61. KODZeIltr.. 525 aud pl. Atti del Coaer. Ii. in OcWmeles a composite and continuous ring of sclerenchyma. CACTEAE. -er. DATISCEAE. and 3 Tab. as is well known.. Anat. Kalkoulattuch..-Peazig. and pI. Petiole.-ZaIewski. SJ)a1tOft'n. n. bot. Schmc:lt:. d. Except in the species of the genus Pereski4 which have thick and fleshy leaves. Centralbl. Digitized by Google . especially in the leaf of OQomeles.I were available for investigation. iii. 240 and Tab. bot. d. iii. iii. are found in the pith. 149-51. z. Bot. have a quadriseriate stalk and a rather large. Ver. and branched sclerenchymatous cells m the pith and primary cortex. Milnch. In OQomeles sumatrana a one-layered hypoderm is developed beneath the upper epidermis. Gaston N~ 100 pp. Obs. p. the cells of the shield have an approximately quadrangular shape in surface-view. bot. In the structure of the wood the two species investipted agree in the possession (a) of vessels with rather wide lumina and honzontal.-Radlkofer. and are arranged for the most part radially. Oxalate of lime has been observed in the form of small prismatic or needleshaped crystals. the structure of which is adapted to a desert-climate. (b) of wood-prosenchyma with wide lumina and simple pits. LianeD..-StahI. The leaf-structure is bifacial in both species. D. P. and traverse the entire thickness of the leaf. Natmf. 18g2. which only in the case of OQomeles sumatrana occur on both sides (though in small numbers on the upper side) are surrounded by several epidermal cells. 188g. the scale-hairs.. 1Il00. found in these regions. Org. p. 18g3. L.. The stomata. 137 et seq. Mig. and OQomeles sumatrana. 5540 555. show transitions to simple pitting in Datisca.. Syst. Mem. the following points should be mentioned. foliage leaves of the ordinary form are wanting. p~:8. The vessel-walls. which are approximately H-shaped. and (c) of narrow medullary rays. lard. pp. 18g2.. v. aud 2 pl.-Ber. Bel.. Zeit. B. one-layered. GaellIch. appears immediately after the two normally developed cotyledons. in Sitz. 5. where they are in contact with parenchyma of the medullary rays. GefUsb. Beitr. whilst some of their arms penetrate as far as the cuticle on both sides. di tieDoft.-Fellerer. Bunte LaubbL. Pftanzenfam. P. In Datisca the pericycle contains isolated bands of bast-fibres with wide lumina..

hne MOwn that in PwulitJ the pair of rnud-ceU. since all the investigations hitherto carried outhave shown that many an anatomical feature can be employed in the grouping and diagnosis of the genera and species. CnYW ~". BP. which fall off at an early date. especially to the physiologist. who can readily correlate the structural features with their physiological causes in an Order of plants which (like the Cacteae) is adapted to quite special and uniform conditions of existence. The occasional occurrence of a secondary division-wall. some of which are transformed into thorns. present many points of interest. placed at right angles or obliquely to the pore in the one or the other subsidiary cell (as Benecke and others have remarked).are alsoof interest to the systematist. The anatomical features of the Order. which will be described in the following pages from statements in the literature. The stomata are commonly found at the base of saucer-like depressions on the I DeveJopmeotal investigation. this sometimes occurs in species of Pereskia and M ammUlaria. subsidiary cella arise by sabdiYision of the initial c:e1l (Strubar&er). M~ ~. according to Caspari. II. is of course not to be regarded as an exception to the general type. The stomata are found in large numbers on the shoots FIG. the anatomical features.CACTEAE and only bears very small foliar structures. and the Digitized by Google .however. also bear stomata.denniaand Rl'IIOderm ofllOlDeCaeteae: of the Cacteae and also on the leaves A. go. however. The guard-cells are accompanied on both sides by one or more subsidiary cells. The small Mter Scbleiden. parallel to the pore 1. and as bas been confirmed by all the later authors. The stomatal apparatus is characteristic of the entire Order. the same applies to the complete suppression of the subsidiary cells on one side of the guard-cells. as Schleiden first pointed out. the latter. are abortive or anomalously developed. rudimentary leaves in Of>tmIia.~ in the species of Pereskia.

The direction of the pore of the guard-cells with reference to the longitudinal axis of the shoot is very noteworthy. e. or Cereus pruinosus. R. floccosa. it may be pointed out that Vochting describes a one-layered hypoderm. according to Schumann).one-layered and collenchymatous in EpipA'YU"m. and is mostly oollenchymatous. EpiphyUum and Opuw according to Lauterbach) it is parallel to the longitudinal axis of the shoot. Dyck. go. and is consequently horizontal j in another group of genera (Lepismium according to VOchting. In connexion with Schleiden's statements regarding the hypoderm of the Cacteae.' solitary crystals in Melocactus macrocanthus and Cereus Curlisi. and O. Melocactus macrocanthus. The cuticle is strengthened by the formation of cuticular ridges (EcJKnocactus according to Lauterbach) and thickened by a deposition of wax (in the form of a covering. Dyck or Qpuntia.CACTEAE surface of the shoot. R. R.) or in the furrows (R. S. two-layered in PA'Yilo&/ICtus. often branched pit-canals. Dyck. and appears to be characteristic in certain genera. brasiliensis. Cereus serpentiftUS and C. 2-3Iayere<l and collenchymatousin Ec"itwpsis. a feature evidently of advantage to desert-plants. S. Dyck. the stomata either occurring on the surface of the ridges (R. The wall of the hypodermal cells is gelatinous and traversed by abundant.or entirely suppressed inM ammillarUJ. The lumen of the collenchymatous hypodermal cells commonly contains crystals: for example clustered crystals. as occurring in PfeitJwa cereiformis. I peruviana. S. the latter in RlJiPsaUs paradoxa. sometimes undulated lateral walls. Phyllocactus and Peres"Ma according to Lauterbach). according to VOchting). Cassytha. have sometimes straight. s. Dyck. and must therefore be referred to the cortical tissue. which shows stronger thickenings at the comers. and is thus vertical. in Pilocereus HouUetii. varies in thickness j the former is lUten continued from the pore of the guard-cells over the entire wall of the respiratory cavity (Mohl). conferta.. A-B). Vacht. which radiate out from the small lumen of the hypodermal cells and are sometimes enlarged into a narrow border at their extremity. grandifiqrus). somewhat thickened h)'poderm in Lepismium.flera and RhiPsalis according to Yachting. showing varied distribution in Qpuw monacamha. S. /unaUs.ch. According to Schleiden. 1-2 layered and slightly collenchymatous in RAipsalis. and a two-. which have to diminish transpiration as far as possible. which is either loose and readily rubbed off or firm and continuous. as also the entire external wall. Digitized by Google . 2-4 layered and strongly collenchymatous in OpuftlN. Thus in some of the genera (P/ei. 2-3 layered. S. It is sometimes sharply marked off from the remaining tissue (Cereus tetragmus and C. Dyck) j and (b) thelappear8. The epidermal cells of the shoots of the Cacteae. The following special structural features of the epidermis remain to be mentioned : (a) the formation of furrows and ridges on the epidermis (species of Rhipsalis of the group Teretes. which consists of one or more layers.g. Echinopsis and Cereus according to Lauterbach) the pore is placed at right angles to the longitudinal axis of the shoot. the former. According toLau terba. and VOchting records it in a few species of RhiPsalis.oides). it does not originate from the dermatogen.llce of relatively thin secondary division-walls in the epidermal cells (species of RhipsaUs according to Vachting. Gaertn. a one-layered. but less collenchymatous in Cereus. sometimes not very distinctly delimited. pendula. The cuticle. Beneath the epidermis there is a hypoderm (Fig. the hypoderm either consists of a single row of cells (in Cereus Curlisi and yrobably in all Mammillarias) or of two rows of cells (Cereus variabUis) or 0 three or more (Qpunila monacantha. PhyUantlt. while in a third group (MammiUaria Echinocactus and Phyllocactus according to Lauterbach) the pore is irregularly orientated. the hypoderm is one-layered and collenchymatous. Lem. EcJKnopsis. 2-5 layered and collenchymatous in Ec"iMCfJClus. ECMnocactus Eyriesii. rarely one-layered hypoderm in the species of Rlupsalis. Caspari mentions this as being quite general. O. as Schleiden first noticed.

resemble R. but also around the normal ring of bundles. in a few species of Cereus with thick shoots (e. Dyck. according to VOchting. in groups of two or more.). DC. Pflf'fIdoxfI). In many species of Rhipsalis (for details see VOchting) parenchymatous cells with thick walls are included in the ground-tissue. fiOCCOSfl. Mammillaria angularis. owing to numerous anastomoses. candicans ?).): the cortical bundles here lie in small H groupe. Pfr. has been described in detail and-ligured by Van Breda de Haan. mesembryflnthoides. In the neighbourhood of the epidermis and in the primary medullary rays the ground-tissue consists of more closely packed cells. and R. The bundles of the normal vascular ring are concerned to a very varied extent in the formation of the axis. and only in so far as the cortical bundles occur not only in the wings. A transverse section through the shoot of Rhipsalis f'Aombefl. micf'flfltAfI. Saglionis. aw. S. they Digitized by Google . medullary bundles are only present in a few Mammillarias (e. VOcht. in the wings of all these plants the cortical vascular bundles lie in two rows. pm".except that the outline then becomes three-. Pfr. shows a narrow band-sha~ outline. and according to K. In correspondence with the different degrees of development of the vascular bundles the composition of their xylem varies..Yacht.' 1l.CACTEAE but absent in Pereskifl.. which contains crystals. pmdulfl.. R. CassytAfl. f'hombefl. They form a dense network in the longitudinal direction. conI""'. RhipsaUs. luna/is may serve as the type of the species 01 RAipsalis belon_ging to the section Teretes (R. According to Mohl and De Bary. M. in addition medullary or cortical bundles occur in certain Cacteae. in some shoots of • Lepismium. R. Otto. SWflMaM. quite in the normal way. Dyck. The cell-contents of the ground-tissue will be referred to below. f'adicflftS' and all shoots of Upismium K fligAtii and PleiQera cerei/ormis. Dyck. C. Dvck. Haw. also found in the portions alternating with the wings. S. the bast-portions being directed towards the exterior. See also Michaelis' statements regarding this feature. belonging to the sections Alatae and Angulosae (R. but not in M. R. and consists (in Ofruntia monacantlta and Echinocactus Eyriesii) of alternating layers of cork-cells with thin and thick walls. LePismiutn and Pfei. according to Yachting) and the forms of LePismium. according to H. in Echinocactus. PflCA'YPterfl. and R.tisaue of the stem is principally composed of ellipsoidal cells. in which the woody tissue of the vascular bundles forms the chief portion of the axis. so far as is known. Amongst the Alatae. PusUla. Cork. R. or many-angled. paf'fIdoxa. saliCOf'1Joides.fJera. a R.. the central portion being occupied by the normal vascular ring. When more than two wings are present. and show an enormous development of the ground-tissue. R.g. while numerous cortical bundles. and form small rings of bundles among themselves.. R. Schumann also in Phyltocactus. Vacht. Pfr.formation. R. the position of the bundles remains the same as in shoots with two WIngs. crisPalfl. cortical bundles occur in species of RhipsaUs. Pfr. all transitions are found to those species which possess weakly developed vascular bundles. In Pereskia.. S. Gaertn. is superficial in this Order. four-. Cflf'nOSfl mongst the Alatae. R. The vascular bundles are in general arranged in a ring. forms the sole exception. Schenck. lunalis. as is the case in many of the segments of the shoot in ~ecies of RAipsalis. Beginning with Pe1'eskia. The ground... they are essentially distinguished from the latter by the fact that prosenchymatous cells never occur in them. In these cases cortical bundles are sometimes (R.pterfl. Pfr. and some Opuntias the compact wood consists of (a) wood-fibres with thick walls and simple slit-shaped pits. and thus grow in thickness like the bundles of the normal ring. most species of Ce1'eus. S. characterized by their sharp angles. Lageftflria. Almost all the Alatae in RAipsalis (R. which are separated by intercellular spaces of corresponding dimensions. are found in the two wmgs. The cortical vascular bundles have a cambium. as in R. and according to Lauterbach also in ECMnopsis.. the cork originates in the epidermis or subepidermally.g. glochidiata and others). According to Schleiden. R. The strongly developed collenchymatous hypoderm of MelOCflClus. RAipsalis Cflf'nOSfl.

In other species of Opuntia they appear sparingly in the articulations between the segments. as in Opuntia tunicata. which are inserted on the thin wail by their narrow edge. <lc. (c) wood-parenchyma. Poniono(thn~of&A"'" Lauterbach). which accompanies the vessels. the vessels attain a diameter of ·23 to ·25 mm. they are generally described as tracheides. and o. and finally of the excretory and secretory structures. According to Schleiden. who demonstrated a protoplasmic sac and a nucleus in their contents. Plum. 91) occur in the wood. opuMa. Finally. rhoda"'lUJ.410 CACTEAE are occasionally septate in Pereskia aculeata. arranged in a ring. laria. (b) pitted vessels. g. aflllieola. O. and PhyUocactus. in Mammillaria simplu. which have been found by Delpino and others in some Cacteae (species of Cereus. The extranuptial neetaries. and Rltipsalis). Schenck. they are provided with local annular or spiral thickening ridges. in other cases they themselves become an integral part of the vascular bundles. the cells are sometimes provided with pits or with superficial striation Glandular hairs have not been observed in the Order. <lc. are found in the primary cortex of Lepismium sarmentostmJ. either 10 the tissue of the medullary rays lying between the bundles. M ammi. however. in Pereskia aculeata. in Echinocactus. a1ldicola.all other elements. and in form are midway between the shape of a barrel and that of a spindle. according to Michaelis). which are sometimes accompanied by spirally thickened vessels in the middle of the secondary wood. in which the individual elements are connected by means of perforations. or at the margin of the pith. 'sWligera.u. biseriate or multiseriate. monacantiJa.-After: Sc:hleiden. O. They are found in still larger quantities in Echinocactus and Muocactru. tiIIdrII E". The hairs are (rarely) unicellular (Echinocaclus. In the genus Opuntia they are also absent in some of the species. The bast of the vascular bundles is strengthened by hard bast in certain members of the Order (species of Pel'eskia. with the exception of the tracheae of the primary xylem. O. the multicellular trichomes are composed of cells which are short and narrow in the basal portion of the hair. then of the thorns.ria and M elocactus is 9uite different. (which is a high climber). According to Caspari. According to H. Dpuntia. where-except in the lowest internodes-they already form the most important part of the xylem. especially in all those which have flat oval shoots of the type of Dpuntia Ficus i1ldica. imbricata. and project far into the cell-lumen. In the following paragraphs I shall speak firstly of the hairy covering. In Echinocactus and Mam". and in no way enter into the vascular bundles themselves. (VOchting). 91. Similar groups of hard bast.-iuii. e.' <le. rows of tracheides occur. the structure of the wood in most of the <>puMas.. in Dpuntia cyli1ldrica. speaks of them as parenchymatous cells. uniseriate. Voecht. Generally these tracheides are also dis~hed from the actual vessels by lack of perforations. disappear and the peculiar tracheides form the bulk of the xylem. M. e. ramuli/era. O. according to SchIeiden and FIG. but Van Tieghem.lltl.M. These cells seem to be entirely wanting in the species of Pereskia and RMpsalis. Cereus. In these cases elementary organs of quite a special nature (FIg. RltifJsalis. and also in the stem of the species of Cereus. brasiliensis. Mammillaria. Plum. have nothing to do with Digitized by Google . and have simple perforations. whilst they are broader and longer towards its termination. They are broad cells with very thin walls. g. and (d) lignified medullary rays..

Schumann. Melocactus. grandifolia. This is not the place for a detailed discussion of the reasons for regarding these structures as foliar organs. Eclainopsis.0f!I::: chymatous. Wetterwald. As regards the structure of the thorns the following general statements may be added. imp.' which perhaps belongs to Cereus). They are arranged on the thorn in longitudinal rows with a spiral course (from right to left in the botanical sense). 92). It must suffice to refer to the interpretation of Delbrouck. Goebel. In the latter ~~ a difierence in the structure of the apex and base of the thorn is also (rom the noticeable. Foerst. Plum. Haw. alODe c:ootain a YUCalar bandle ill their bual portion. while those of the base are mainly paren. Anhalonium. and RAipsalis).). in the two species of the group Subcarnosae. arising in the same way as the thorns. =~ Digitized by Google . which is more or less distinctly produced into three points. whilst the apex of each cell is covered by the freely projecting end of the preceding cell. Foerst. spatlaulata. has been most thoroughly examined by Caspari. Pereskia. PiloceJ'eus. and P. uDV.' to the views of Caspari and K. the determination of the morphological value of which has occupied a number of investigators. Pfeiflera. Bk<J. e. and invariably show a difierent structure at the apex from that at "the base of the thorn. Their epidermal cells invariably possess straight and never undulated lateral walls. RMpsalis. oripaL " EcMMcactUS. and immediately surrounded by pitted sclerenchyma-cells with relatively thin walls. Link. According to Caspari. Kauffmann. subulata. P. The I The traDlitiOllal forms between thorns and lea_' which KaufrDWlll fillt obeenecl forty years ago (BulL Soc. but are cushion-shaped bodies. and Dp-unIia. who consider the structures in question as being of the nature of emergences. but not in the species of the section Frutescentes. Vascular bundles generally take no part in the formation of the thorns 1.. but is due to the fact that the cells of the apex are cuh~ rA chiefly prosenchymatous. and serve as hooks. This difference is most clearly marked in the thorns of all the species of Dpuntia (with the single exception of • opuntia clavarioides cristata. The structure of the thorns just described had previously been pointed out by E~elmann as highly characteristic of the genus Dpuntia. EclaiMcactus. In the Opuntias (see Fig. and to the works of Zuccarini. 92) and the two first-named species of Pereskia the integumental cells of the tip of the thorn are scaly. and P. aculeala. and excreting a fluid containing sugar. The latter consist of a bundfe of sclerenchymatous cells with thick walls and devoid of pits. they are flat and lie in an imbricate manner over one another. The free ends of the scale-cells are directed backwards towards the base of the thorn. which should perhaps be united with Opuntia. Cereus.E epidermal structures. de Moscoa. who regards the thorns of the Cacteae as a transition between' leaf-prickles' (Phyllomstacheln) and ' trichome-prickles. de Nat. Lepismium. P. His observations extend to more than a hundred species of the genera Mammillaria. Cereus. the last-mentioned features can only be employed in specific diagnosis. The outer covering is composed of epidermal cells. PfeifJera. and in the thorns of some Pereskias (viz. In addition to this the basal cells sometimes exhibit markings or prominences on their membrane (as in Mammillaria. which have wide lumina. PiloceJ'eus. Miihlenpf. and as fornung a point of difierence from the other members of the Order..CACTEA. who support the theory of the foliar nature of the thorns. DC. with P.. The structure of the thorns (Fig. 1859) and which have more recently again been met with by Gmoog.: i. and Ganong. which will be described unmediately.

or from the cell-membrane (according to De Bary and Walliczek). Linke. gracilss. M. M.. pukAella. SOmetimes crystals occur in the mucilage-cells. subsetostU (M.. Pilocereus.. subecAifUJla.. in most cases.. DC. Haageau. julvispiu.. Zuccariniana. anpinea! 0to. elongata. S. gladiata. Lem. Pfr. but laticiferous canals and mucilage-cells are also found. Rhipsalis. M. angularis. M. Cweus. M. decumana. Dyck). The mucilage-cells are found in the whole of the ground-tissue. EcAinopsis. owing to a single solitary or clustered crystal occurring in each cell of the bypoderm: in Pelecyphora. Echinocereus. tumcata. M. MOfIvillei.. or from both (according to Longo ?). M. whilst an irregularly shaped mass of mucilage usually occupies the middle of the cell.. stelligwtU (M. Berol. crocidatlJ. MammillaritU IcmgimammtU (M. M. and a few other cases of rarer occurrence may be added. glocAidiata. Scheidw.bergiapriflCipis. Echinocactus Oltonis... M. under the protection of which the cells of the thorn gradually undergo sclerosis. M.). M. M. subangularis.. Mart. Ehrenb. Link et Otto. De BUy. macracaflthll. these canals are situated extemally to the bast of the more strongly developed vascular bundles (Schleiden. Haw. (b) forming a connected layer beneath the epidermis. M.412 CACTEAE angle of inclination of this spiral line with reference to the longitudinal axis of the thorn sometimes varies in the same thorn. Mart. Berol.. M. M. Of these the crystalline excretions of oxalate of lime are the most important. rAodaflthll.) consists of thick-walled cells. M. DC. In addition to the mucilage-cells lysigenous mucilage-canals occur in a number of species of Opuntia with the segments of their shoots flattened like leaves. also Leuc/Jtnl. RAipsalis paradoxa.Engelm. a1lgulares(M. Dyck. Poselg.). (b) crystal-cells forming a connected layer beneath the epidermis: MammiUaria macrothile. S. M..). Dyck. Bockii. Fowsteri. Mammillaria 1Iiera. Foerst. cmtrici"hII. Lem. &c. lcmgimamma. M. Lem. M. elegans. and E. Crystal-cells alone are present: (a) irregularly distributed in: AnIJaloniulfl. fJlaCf'omens. and Longo 1). Mart. Mart. Opufltia. (here the laticiferous canals have scanty contents). rhllpAidacaflthll."a. running parallel to the cell-wall. M. Aystrix. M. and Pweskia. It has not yet been definitely determined whether the contents of the mucilage-cells are derived from the protoplasm (according to Lauterbach). viridis. M. Astf'DPhytum. hsteracamAae. The laticiferous canals. M. M. Haw. Miihlenpf. Lem. Lem. M. and is therefore not of systematic importance.).. Engelm. DC. M.... Mart. Melocactus. and Echi1fOCactUS. PfeiUwa. Before entering into details regarding these points we may prefix the synopsis given by Lauterbach of the occurrence and distribution of the various structures in question : I. EPiphyllum. doIic/Jo". pro parte (M. Mart. ~AocepAala. Bocasau. are found in many.. M. as is apparent from the above conspectus.. M. Dyck. M. Miihlenpf. ci"hitwa. M. II. Dyck. The excretory and secretory structures of the Cacteae have been studied in special detail by Lauterbach. S. DC. Pfr. although he lell%Ched for them.. and M.S. Lepismium. sanguinea. Miihlenpf. M. Webbiau. Krameri. Lem. g(auca. Mart.. Lem. S. magnimamma. applafUJla. SIella-aUf'ata. PwesAia subuiata. Emufllftiiau. crinitae (M. DC. III. megacafltAa... M..)... Mucl1age-cells and crystal-cells are present: (a) crystal-cells irregularly distributed: MalacocarPU5.. but not in all species of MammUlaria. Lem. their physiological importance as water-reservoirs is evident. as well as in LeucIttMI Lauterbach curiously enough did not oblene theIe c:ana1s. Haage.. HoPtwiau. Mammillaria elephantidens. M.. The following possess laticiferous canals and crystal-cells: M ammillariae centrispintU (M.).. pentacafltha. Lem. The coloration of the thorns is caused by the deposition of colouring-matter in the membrane or lumen of the inner sclerenchymatous cells. Hort. multiceps. Neumanniana. M. Phyllocactus. ""tocrocea. M. Dyck. M... Digitized by Google . The calyptra of the prickles of some Opuntias (0. Dietr. Haw. S. The marginal portion of the mucilage exhibits stratification. Link et Otto. Hort. Hort.. such as O. M.. Fisch. M. Pfr. M.

and send numerous branches into the superficial. SchUDWlll. on the other bud. A so that it forms as much as 85 per cent. likewise consisting of monoclinic prisms. and may therefore be considered as a mixture of resin. In this genus their contents consist of a thick whitish sap. of the weight of the ash of the entire plant (in Cephalocereus senilis. but they do not appear to extend into the small lateral roots. Oxalate of lime is sometimes excreted in enormous quantities in this Order.CACTEAE wcia principis 1. chlorophyll-containing parenchyma.14 mm. the most widely distributed form is that of star-shaped clustered crystals. Gutwinski in his paper points out the fact that the laticiferous canals of Mammillaria are never met with in the pith and medullary rays. these contents are insoluble in water.14 mm. tetragonal and monoclinic prisms are found: Bundles of raphides. According to Lauterbach. and E. but soluble in chloroform after long exposure to this reagent. and the latter vary in shape. Solitary crystals are rare in this Order. composed of monoclinic prisms with a very long principal axis. They are to a certain extent characteristic of groups of species in Mammillaria. accordiDg to Michaelis. The mode of origin of the laticiferous canals is lysigenous. and hence occur in the cortical parenchyma only.. Both solitary and clustered crystals occur. especially into that of the warts.' CODtrary to the CllItomary DomeDciature. In the cortical parenchyma they anastomose with one another. but they increase in abundance towards the periphery. laticiferouscaDW are Dot loud in thele tW9 species. which. and also soluble in ether. composed of monoclinic prisms.and caoutchouclike substances. In the cortical parenchyma of the roots we again meet with these canals (with a diameter . according to Schleiden). who describes the _tory 01_ . quadrate octahedra. which rapidly hardens on exposure to the air.). they are especially characteristic of the Dpumias. The diameter of the canals is between '042 and . however. also poMeII latldferou canals according to K. Digitized by Google . '<>7-. and Incorrectly ucribel a &ehizogeDou mode of origin to them (see below).. according to Gutwinski and Lauterbach.. In the interior of the plant the canals are isolated. in question u 'latex-II&CI. LIwi"u. HeDII. have been observed by I &lritUKlKllU WiOiamsii. where the canals extend to the region just beneath the hypoderm. they are stained bright red by tincture of alkanna. Lem. have quite a short principal axis. Almost as abundant as these' are very irregular clustered crystals. With regard to the distribution of the laticiferous canals the following facts may be pointed out on the authority of Lauterbach.

MIlL d'hist. To complete the discussion of the cell-contents in the ground-tissue of the Cacteae. u. u7. h. p. _Ilea et apiralc!es des C.. Cadau-StachelD.. Man. GeseJJsch. Literatale: De CaDdoUe. Acado des leo de St-P~ Ih. rril.--&bJeideD. ~". Cell. ZD Lemberg. CentralbL J887. which is sometimes rolled up. 369 and 409 et seq. t. P. about which the outer layers are arranged. island-like areas. ProteIuki:irper in den Zweigen von Ejlil"~U-. Wiener AbeL 187. 1828. the first two forms are commonly stratified to such an extent that they appear to be composed of threads. Do :a-a. with Latin abIItr. imp. Pringsheim Jahrb. in T~ ~. lying in intercellular spaces. 1885. DXi-D. DiII. According to Kohl.. SeIaetbeh. pp.-Mobius. 6. and 1 Tab. p.-Laaterbach. Molisch regards these structures as reserve-substances.CACTEAE Lauterbach in OPUnlia Ficus indica and EcMnocactus OUo1&is. bei den C. iii. Finally. PB. Cact.. VergL Anato. Crystals of oxalate of lime are excreted in the cell-wall of the epidermis and of the subepidermal cell-layer in Pereskia Bleo..Moller.-Delpmo. 8. with c. Pff. 1883..tecb.f8 et seq.-Caapari.-Miquel.-Vi)chting. remain to be mentioned. d. 195 et eeq. Bot. Soc.-W • Anat.eo.. deutIch. <_ p"~n_. bot. Bel. it is doubtful whether these sphaerites are of systematic value for the species in which they occur. Digitized by Google . d. RhiJ)l&lideen. I03-6.Gutwinski. SphiiroIaJItalle.. Halle a. Anat. Merkw.. 1845.-Strubutger. and in the root of Cereus rostratus. Soc. 1839. 1890). Ber. and to the place of occurrence of the latter. 1845. d.. without however entering into the special structure of the sphaerites in the different species: thus sphaerites are found in the apices of the mammillae of Mammillaria W ildiana and M. and often attain considerable length.vlii. 1873-4. and 10 Tab. P. Anat. it remains to mention the frequent occurrence of sphaerocrystaIline masses of undetermined nature. natmaL de MOIICOIll859. It is also necessary to say a few words on the curiously-shaped proteid-bodies (Fig. Botanical Works. 1885. they have very thin walls and on superficial observation might be taken for clustered crystals.. 181-242. 1875. 1. 71-6. 93. Entwicld.-De1brouck. U pp. Bel. Cereus rostratus. v. t. d. p. C. In all species of this genus large proteid-bodies are found in every cell of numerous. separate copy £rom Jahrelber. pp. Chmielewsky. i. Cact. according to the same author. LiImaea 1842. Adansonia. deutach. d. 205 Dote. DaL. MudJageI. 'Bd.eq. Tijcbchrift voor D&tn1IrL GeIcIUed. iv. in the stem of EpiphyUum sp. pp. de FJ'IJDCe1885. their diameter varying between '0125 and ·02 nun. 2. SpaltOBu. Bot. 28!).. FllllZione mbmec:ofila net regno T&b. beloaging to the Haemadoraceae (Wakker. 1887. nycticalus and PhyUocactus grandis.. as they contain several centres.S:-V' 1 Similar bodies are found in the On:hicl 0IIei4iIIIII (MikOlCh. 46 pp.. etc.-Euaelmann.f65 et . Ber. 93..-VUl TJeghem. which contain clustered crystals and have been described by Kasimir. Anat. t. BaD DDd EDnr. Franz·J __ Gymnu. in Venn.-TRcaJ.... which occur in the epidermis and the neighbouring cortical parenchyma. d.. DeDbchr. vii. Sometimes they resemble half-compound starch-grains. BoloenalJ:!. and detailed IItateDJeDts about it). the threads are found either singly or several in the same cell. Polish.. Nevertheless brief reference may be made to a few plants containing sphaero-crystals. P. in Prinpheim Jahrb.n. KauftilWm. 257.6. Morph. ProteInk.. Dia. bot. HohaDat. {'po 74-5 and 37o. PringsheinJ Jahrb. CentralbL 1889. des Man.-De Buy. which Lauterbach and others have found deposited in alcohol material. Schrlft. pp. 1877 hae the older literatare... 32!). 1888..-Stach. on the basis of Kohl's observations. deutsch. IWustein. thin thread. 1~7' po 317. 3'7484 and Tab. Ball. They have a spherical or ellipsoidal shape.-GeIdL d. Ber. pp. bot. GeIelIlc:li. very fine sphaero-crystals (Fig. A-B) of oxalate of lime occur in some species. Milchginge bel der Gatt.-Mohl. the idioblasts of Dpunlia Lemaireana. Bel.. 178 ~1Cb Molilch. and l30L Zeit. BeL iii. Fun. . ChmieJewsky as secretions J.ie1ocaeten. bOt.1#". Abh •• ii. a ring or a very long. po 3. which are united by a substance of a different refractive index. gee. etc. _ EIt"Bot. C-E) which were discovered by Molisch in the branches of EpiphyUum. in put Dot CIted JD the preIeI1t review. C.-Nik. 1888.HartiJJ2. 1866-7.. Ball. nUi) and in the teDtac:Ja of DI'fJII1'8 tlicllot_ (Gardiner). en Ph. According to Moebius and Kohl. iii. P• . As a rule these bodies have the shape of a straight or bent spindle. 115 Dote. Hantgew. BaD d. M_illarUJ. 1842. macrotlsele. d.

-Walliczelc. Meyen. Auuuar. della Soc. R. d. Dannemann. Membranschlelm. iii. baDd.. ftII het ~t Md«tIttlU. in which the bladders on the upper side of the leaf suddenly narrow down at the apex into a short and slender hair-like structure.--GaDODg.g. The anomalous structure of the axis is. De Candolle. ~ 5~Lougo. 111930p. Anat. embedded in a prosenchymatous or partly parenchymatous ground-tissue (Mesembryanthemum-type). internal secretory organs are wanting. Herbler Boluier. Miinc:beu. Morph. d. Schenck. Ball. 123 pp. 14 pp. u.. and "lilora 1119. The former must be regarded from a physiological point of view as water-reservoirs. Disa. bladder-like epidermal cells. bot. the lack of parenchymatous medullary rays. 6 pp. 940 A-B) and of aqueous tissue in the interior of the leaf m many species may be especially pointed out. has been specially pointed out in many members of the Order by the various observers (Schrank. the simple pitting of the prosenchymatous groundtissue. 262 et aeq. water-storing.. bot. subsidiary cells are present.CACTEAE Tab.. Fig. iii. Kiesel . FICOIDEAE. (Fig. loe. Only a relatively small number of species O has been investigated with regard to their leaf-structure.~UIlWlll.)-Michaelis. Cork-formation is (at least in MesembryanthemtmJ) internal..-Lougo. which may be concentrically or irregularly arranged.. bladder-like cells in the e:pidermis (Fig. Gatt.-H.. Bd. ii.. Anat. vii. and 3 Tab. 11196. 940 A-B). 1889LP. and 1 Tab. where the bladder-like cells are attenuated in a papillose manner at the apex. and stellate hairs with a uniseriate stalk and a terminal cell. Tell. ii. The highly characteristic epidermal differentiation (Fig. 940 A). Mirbel.. 11193. d. . 11193. Large. 2. In the leaf-structure. 111940pp. 6a. ell Roma. 39 pp. A constant and characteristic type of stoma is not found. u. According to the existing investigations the following features in the structure of the axis may be regarded as characteristic: the anomalous stem-structure.. Ital. When completely filled with water they give a peculiar appearance to the plants of this order. 11197. 164 et aeq. however. styloids (axis of Aizoon and Tetragonia) and clustered crystals. p. Heinricher. 148. Hagen.-Van Breda de Haan. 8. unicellular two-armed trichomes (A izoon. which is often found on the stem as well.-Kaaimir. in NatiirL PftIDmd"am. Martinet. Muc:ilaelue delle Cactee. m some of the species of Mesembryanthemum. d. assuming the form of a star (Glinus and MoUugo respectively. REVIEWOF THE ANATOMICAL FEATURES. In addition to the two Digitized by Google . are found intercalated among the cells of ordinary dimensions. only present in the species with thick stems (exception: Polpoda). Zeit. xxiii. and the simple perforations of the vessels. found principally in the outer wall of the epidermal cells of the leaf. and Volkens). MaJII"'. 1st. The hairy covering consists of simple unicellular hairs. and in Mesembryanlilemum crystallin""" L. however. &"itf«adus. Oxalate of lime is excreted in the form of raphides (Mesembryanthemum and Gisekia). Diss.-Kobl. Bot. in these the transverse section of the axis either shows several concentric rings of vascular bundles (Tetragoniatype) or bundles. to which the latter owe the name of Ice-plants.-lIeDecke. Ielloblastl muciferi delle C. and 1 Tab. the occurrence of large. ErgiiDzaDgI. 11196. cit. e.. AdaItmi_ etc. t. C. and later by De Bary. 32. which are placed parallel to the pore. Biol d. Llanen. PP' 5G-2. 586-8. Kalks. With the exception of very large tanninsacs occurring in the leaf-tissue of some species of Mesembryanthemum. 1891. Abt. vol. and 3 Tab. which shows a distinct adaptation to the dry desert-regions where the members of this Order occur. sometimes reaching the size of peas. pp. Nebenz. 940 C). and Iutomo ai c:ana1i delle OpUDZie. I. in Tetragonia expansa. Some of these large cells occasionally tend to form hairs. Harlem. and of the wood-prosenchyma. Bull. SpaltOfl'n. STRUCTURE F THELEAF.. and Plin"'us). PriD2sheim Jabrb..l/ivia u. in Mesemlwyanthemum it also occurs as a deposition in the cell-wall in the form of small crystalline granules. Galenia. Er1angen and Halle a. 111940p. VergL Aut. Criltaux chez Opull/ill et PWeSM.