THE NEW PHYTOIiOGIST.

VOL. 4. No. 7. JULY 31 ST, 1905.

PROVISIONAL SCHEME OF THE NATURAL (PHYLOGENETIC) SYSTEM OF FLOWERING . BY HANS HALLIER, PH.D. ().

S the son of a well known botanist and the grand-nephew of M. J. Schleiden, the centenary of whose birth was celebrated lasAt year at Hamburg and Jena, I have by inheritance and edu- cation a passion for , and for nearly thirty years 1 have taken a zealous interest in . Even at the age of seven to thirteen years I used to accompany my father, the late Ernst Hallier, formerly professor of botany at the , on the botanical excursions which he undertook every Saturday or Sunday with his students, in order to introduce them to the principles of systematic botany. In these instructive excursions I soon gained a knowledge of the botanical names of plants and of our native flora, and learned the natural system by immediate intuition and comparison of the objects themselves, independent of the sometimes very dogmatic views of the standard-books, or, as we say in Germany, " mit einem noch nicht durch Fachkenntnis getriibten Blick." Later, at the University of Jena, I was intro- duced by the works of Darwin and the lectures and practical instructions of Ernst Haeckel, to the marvellous series of dis- coveries, to which the evolution-theory had been the impulse. In the Botanical Laboratory of Professor L. Radlkofer and Dr. H. Solereder at Munich, I recognized that not only the external characters of plants must be examined in determining their affini- ties, but that comparative anatomy is also indispensable to systematic botany.' During a four years' stay at the famous botanic garden of Buitenzorg and during an expedition into the

' See H. Solereder's Systematische Anatomie der Dicntyledonen. , 1899. 152 Hans Hallier. centre of Borneo' I had an excellent opportunity of examining representatives of nearly all the natural orders of Flowering Plants under natural conditions. The academical lectures of Professor Ernst Stahl, as well as the personal influence of this suggestive botanist and of Professor Jean Massart, my enthusiastic companion in many excursions in tbe vicinity of Buitenzorg-, induced me to apply to systematic botany, the results of -ecology and mor- phogeny. As a result of these various influences I reached the conviction that there can exist only one really natural system, namely that which is identical with the tree of descent; to recon- struct this, systematic botany should be founded on a much broader and more universal base than at present, comprehending not only the morphology of the reproductive organs, but also all the other branches of botany, such as comparative morphology of the vegetative organs ; comparative anatomy, ontogeny and embry- ology ; phytochemistry, physiology and ecology ; structure of pollen and seed coat; relations to climate, seasons and to the sui'rounding organic world ; plant geogi'aphy ; palaeo-phytology, etc. Thus since the elaboration of my first botanical publication (in 1890-2) till now I have always paid special attention to these points in reference to natural affinity. The results of these, comparative studies have been published in a series of papers, most of which refer not to the whole system of flowering plants, but to single orders or alliances. It was only on my second voyage to tlie tropics, that I felt impelled to publish, in April 1903, a provisional account of my system as a whole. At that time I had to take into account the possibility that the Tagal people might not let me out of the Phillipine Islands, or that a typhoon might throw me into a shark's mouth or into the depths of the South Sea, and I therefore thought it wise to publish my system at once, incomplete as it was. Moreover I hoped, that during my absence competent botanists would have time and leisure to digest my new system and to subject it to discussion, but in this latter point I am somewhat disappointed. Hitherto reviews on my system have been few in number and several of them very diffident or even unfavourable. The more favourable ones are the following. • The results of the botanical explorations of this expedition have been described in the Natiiiirkiiftdig; Tijihchrift voor Nedcr- Inndsch Indie LIV. (1895) pp. 406-449 and in the Naiurwisscn- scliaftUchc IVoclwn sell rift, XI. (Berlin 1896). ' See his interesting descriptions of these trips in his paper, Uu botaiiiste cn Malaisic, published in the litilletin dc Id Societe royale dc botaniquc dc llclgiquc, XXXIV. (1895), part 1. Phylogenetic System of Flowering Plants. 153 In the preface to his German edition of Warming's Manual of Systematic Botany (1902), Moebius refers briefly to my opinion that the Synipcialac arc of potyphyletic orij|in. On pp. 411 and 137 of the sixth, and on pp. 418 and 443 of the seventh edition of Strasburger, Noll, Schenck and Karsten's Text-Book of Botany (Jena, 1904 and 1905), Karsten recapitulates my opinions of the origin of Dicotyledons, Monocotyledons, Choripetnlae, Syinpetalne and Anicntiflorae. In his paper entitled. Die Grundlagen des Hallierschen Angiospermensystems, eine phylogenetische Studie {Bcihcftc z. Dotan. CcntrulUatt, XVII. (1904), pp. 129-156), Gustav Senn, lecturer on Systematic Botany at the University of Basel, undertakes a very careful and critical examination of the principles on which my new system is founded, and in Just's Annual Report on tlie botanical literature of 1903, Fedde reports at great length on my different publications of that year. Only partly favourable are Wettstein's reviews of my paper on morphogeny and phylogeny (Hamburg, 1903) in the Oesterrekliische Dotan. Zcitschrift LIII. (1903) and in the Botanisclie Zeitiing, LXI. (1903), section II. pp. 311-314. Unfortunately, with few exceptions, this eminent hotanist does not indicate precisely nor in detail which of my opinions he considers as inacceptable, and in his remarks on Gnetacesc, I am convinced that his criticism is not well founded. In external characters, as in anatomical structure, the Gnetacea: approach very closely to certain Lorantbaceae, and to Myzodcndriim, and there is much evidence that they belonjj to this cycle of aflinity, if only we presume that the so-called nucellus of Gnetuvi repre- sents not a single ovule, but a placenta with several ovules, as Treub has indicated in some Loranthaceae. The most striking point in the reception of my system is the silence of Professor Engler. I am much disappointed that a scientiBc journal which is a recognized centre for systematic botany and plant-geography has been unable to discuss in extenso a new system, which is exceptional by the complexity of principles applied in it. But in the volumes of the Jahrbucher since 1902, there is no mention of my publications, and the same is the case in the monographs of the " Pflansenreich." In KOhne's Monograph of Lythracese (Oct. 1903) and in Winkler's Monograph of Betalaceae (June 1904) my publications treating the same topics are neglected. Only Buchenau in an appendix to his monograph of Alismataceae refers to Miss E. Sargant's and my own suggestions on the mutual relations of Monocotyledons and Dicotyledons. 154 Hans Hallier.

The general features of my system are the following: The Angiospcrmae are a natural (monophylctic) group, ;ind not a polyphylctic one, as suggested by Engler in Engler and Prantl's Ndtiirlichcn Pflnnzciifamilieu, Nachtrag zu 11.—IV. (1897), pp. 364—369. The Anienlaccae are not to be considered as old types, remaining in a lower state of development, and iillies or descendants of Gymiiospermae, but, on the contrary, as the highest and most reduced types of one of the lines of Dicotyledons. They and all the other lines of Dicotyledons have been developed by reduction in flower and fruit from the Polycarpicae, the latter group being derived immediately (rom Bcnncttitnceae or other extinct Cycadales. In the same manner the Liliijlorae and all the other syncarpous Monocotyledons have been derived by union of the carpels, by reduction in the numher of parts, by epigynous insertion of the perianth, and by other changes in the structure of flower and fruit from the polycarpous Monocotyledons (//f/o^)?r7f), which latter group originated immediately from the polj-carpous Dicotyledons (Poly- carpicae and Raiiales). In the Dicotyledons the Apetalae and Syinf^etnlae arc unnatural groups of polyphyletic origin. Agreeing in some points with mine, and likewise phylogenetic, is the system which Professor C. E. Bessey, of the Nebraska University, published eight years ago in the Botanical Gazette, Vol. XXIV. I do not claim for my system the position of an infallible gospel or of a " Nolimetangere," the latter very significant expression being used by Professor Karsten on p. 443 of the seventh edition of the Bonn Text-Book of Botany (1905). On the contrary, I freely confess that my system gives only an approximate idea of the lines of descent and of the mutual relations of the Flowering Plants; it is only one step in the further progress of phylogenetic botany. But I am sure that this step is not a wrong and useless one, and that it will lead to a broader knowledge of the natural affinities of Flowering Plants. The following is a chronological enumeration of the publications in which I have worked out my system. 1. Versuch einer naturlichen Gliederung der Convolvulaceen auf anatomischer und morphologischer Grundlage. Engler's Botanische yahrhucher, XVI., 4—5 (1893), p. 486. 2. Betrachtungen iiber die Verwandtschaftsbeziehungen der Ampelideen und anderer Pflanzenfamilien. Natuurhmdig Tijdschrift voor Ncderlandsch Indie, LVI., 3 (1893), p. 486. Phylogenetic System of Flowering Plants. 155

3. Die indonesischen Clematideen des Herbariums zu Bui- Aniiales dn jardiii boUuiiquc de Bicitenzorg, XIV, 2 (July, 1897), p. 249. Contains some general remarks on differentiation and reduction. 4. Ueber die Gattung Eryeibe und die biologische Bedeutung der stammburtigen Bliithen und Fruchte. Bulletin de V herbier Boissier, V., 9 (September, 1897), p. 753. A few general remarks. 5. Ueber Kautschuklianen und andere Apocyneen, nebst Bemerkungen iiber Hevea und einem Versuch zur Losung der Nomenklaturfrage. Jahrbuch der Hamburgischen wissenschaft- lichen Anstalten XVII., z. Beiheft (November, 1900), pp. 200—202. 6. Ueber die Verwandtschaftsverhaltnisse der Tubifloren und Ebenalen, den polyphyletiscben Ursprung der Sympetalen und Apetalen und die Anordnung der Angiospermen iiberhaupt. Vorstudien zum Entwurf eines Stammbaums der Bluthenpflanzen. Abbandlungen, herausgeg. vom Naturwissenschaftlichen Verein Hamburg, XII. (June 1901), 112 pp., with a complete index of the plant names. 7. Ueber die Morphogenie, Phylogenie und der Generations- wechsel der Achsenpflanzen. Vorlaufige Mittheilung.—Berichte der Deutschen Botanischen Gesellschaft, XX., (November 26th, 1902), pp. 476—478. 8. Ueber eine Zwischenform zwischen Apfel und Pflaume.—• Verhandlungen des Naturwissenschaftlichen Vereins, Hamburg, 3 series, vol. X. (1902), pp. 8- 19 9. Beitrage zur Morphogenie der Sporophylle und des Trophophylls in Beziehung zur Phylogenie der Kormophyten.— Jahrbuch der Hamburg, Wiss. Anst. XIX, 3, Beiheft (January, 1903), pp. 1—110, with 1 plate. 10. Ueber Hornschuehia Nees und Mosenodendron R. E. Fries, sowie liber einige Verwandtschaftsbezieheungen der Anonaceen.— Beihefte zum Botanischen Centralblatt, XIII., 4 (February, 1903), pp. 361—367. 11. Ueber die Abgrenzung und Verwandtschaft der einzehien Sippen bei der Scrophularineen.—Bull. herb. Boissier, ser. 2, vol. III., 3 (March, 1903), pp. 181—207. 12. Ueber die Verwandtschaftsverhaltnisse bei Engler's Rosalen, Parietalen, Myrtifloren und in anderen Ordnuiigen der DiUotylen.—Abhandl. Naturw. Verein Hamburg, XVIII. (March 1903), 98 pp. 13. Ueber den Umfang, die Gliederung und die Verwandtschaft I5& Hans Hallier.

der Familie der Hamamelidaceen.— Beihefte z. Botan. Centralbl., XIV. (1903), pp. 247—260. 14. VorlauRj»er Entwurf des naturlichen (phylogenetischen) Systems der Bluthenpflanzen.—Bull. herb. Boissier, series 2, vol. Ill, No. 4 (April, 1903), pp. 306—317. 15. Ueber die Gattung Daphniphylliun, ein Uebergangsglied von der Magnoliaceen und Hamamelidaceen zu der Katzchen- bluthlern.—Botanical Magazine, XVIII. (Tokio, June, 1904), 15 pp. 16. Ein zweiter Entwurf des natiirlichen (phylogenetischen) Systems der Bliithenpflanzen. Vorliiufige Mitteilung.—Berichte der deutschen bot. Gesellsch, XXIII., 2 (March 29i;h, 1905) pp. 85—91. 17. Neue Schlaglichter auf das natiirliche System der Dikotyledonen. Phylogenetische Betrachtungen. Gera-Unterm- haus, W. Koehler, July, 1905.—12 pp.

A.—SPOROPHYTES. I. FlI.lCALIiS. II. LYCOPODIALES (near Equisetales and Cycadales, derived from Marattiales). a. IsopORAE. 1. Psilotaceae. 2. Lycopodiaceae. b. HETHROSPORAE (Ligulatae). 3. Selaginellaceae. 4. Isoetaceae. 5. Lepidodendraceae. 6. Bothrodendraceae. 7. Sigillariaceae. III. EQUISETALES (near Lycopodiales and Cycadales, derived from Marattiales). 1. Sphenophyllaceae. 2. Protocalamarieae. 3. Calamarieae. 4. Equisetaceae. B.—SPERMATOPHYTES. a. GYMNOSPERMS {excluding Gnetacea.e). 1. Cycadaceae (derived from Marattiales). 2. Bennettitaceae (derived from 1). 3. Coniferae (including Salishurieae and Taxaceae (derived from Cycadaceae or near Cyca- daceae, Equisetales and Lycopodiales, immediately from Marattiales). b. ANGIOSPERMS. oc DICOTYLEDONS. I. — PoLYCAHPiCAE (containing oil-cells). • MAGNOLINEAE (liypogynous, Eupomatia and Trocbodendrum excepted). 1. Magnoliaceae. a. Drimytomagnolieae (hypothetical group, derived from Bennettitaceae or near them from Cyca- Phylogenetic System of Flowering Plants. 15 7

daceae, and uniting the primary characters of b, c and d), b. Illicieae (including Trochodendrum? and Tetra- centrum? Descendants of a), c. Schizandreae (descen- dants of a or b). d. Magnolieae (descendants of a). 2. Canellaceae (descendants of lb). 3. Anonaceae (including Hornscbuchia; derived from and (for instance, by Anona squamosa) connected with lc). 4. Myristicaceae (descendants or only a tribe of 3). '•''• L.\UR1NEAB (perigynous or epigynous). 5. Calycanthaceae (liescendtmts of lc). 6. Moniiniaccae (including Gomortega ; near 3 and 5, derived from lc). 7. Lauraceae (including Hernandiaceae ; allied to 5 and 6). 11.—RANALES (no oil-cells). 8. Berberidaceae (including Lardizabaleae, Podophylleae, Glaucidium and Hydrastis; near 3, 6 and 7, descended from lc). 9. Menispermaceae (descended from Lardizabaleae, or close to them from lc). 10. Ranunculaceae (descendants of extinct Podophylleae). 11. Nympbaeaceae (origin near Anemonopsis, Trollius, Caltha, Ficaria, Ranunculus, Batrachiuiii and extinct Helleboreae witb acyclic polymeious perianth ; ancestors of Helobiae and of the whole division of Monocotyledons). 12. Ceratopbyllaceae (including Circaeaster ? Descendants of Cabombeae). III.—RHOEADALES (excluding Moringaceae ; no oil-cells). 13. Papaveraceae (including Fumariaceae ; allied by Romneya to Paeonia and descended from Paeonieae or Podo- phylleae). 14 Capparidaceae (including Tovaria! Near 13 descended from 8). 15. Resedaceae (near Tovaria, Pteropetalum, Ritchiea and Crataeva descended from Capparideae). 16. Cruciferac (descendants of 14). IV.—PiPERALES (excluding Lacistemaceae! Containing oil-cells; most of them with stipules ; mostly with perianth). 17. Lactoridaceae (descendants of lb?) 18. Piperaceae (including Saurureae; near 17, descended from lb?) 19. Cbloranthaceae (allied to 18). 20. Myrothamnaceae (allied to 17—19?) V. MALVALES.' 21. Sterculiaceae (including Dombegeae, Triplochiton ! Buet- taerieae 1 and Goupia 1 Near 3 originated from la). ! 22. Papayaceae (descendants of 21). I 23. Eupborbiaceae (excluding Buxaceae I and Daphniphyllum ! Near 22, 24, 28, 29, 117, etc., descended from 21). ' The arranj;cmeiit and limitation of this and the followinj; cohorts is only a provisional one and is very diffieult, because most of them represent parallel lines, having the same point of departure in Stereuliaeeae. 15^^ Hans Hallier. 24. Bombacaceae (descendants of 21). 25. Malvaceae (descendants of 24). 26. Elaeocarpaceae (allied to 21, 24 and 27). 27. Tiliaceae (including Ropalocarpus and Catostemma ; descendants of 21). ! 28. Rhamnaceae (including Neopringlea ? Allied to 21, 23, 27, 29, 62, etc ) ! 29. Urticaceae (including Ulmaceae, Moraceae and Canna- bineae; allied to 23, 27 and 28). ! 30. Dipterocarpaceae (allied to 21, 24, 26 and IV.) VI.^EBENALES (descendants of V.) 31. (allied to 26, 30, 32—35). ! 32. Convolvulaceae (including Cuscuteae, excluding Nolaneae; allied to 31). 33. Ebenaceae (excluding Brachynema; allied to 31 and 35). 34. Symplocaceae (allied to 35). 35. Styracaceae (allied to 30, 31, 33, 34, 54, 80, 101, 111, etc.) VII.—GERANIALES (near 62 - 64, descended from 21). 36. Zygophyllaceae (excluding Peganum ! Allied to 38 and 39?) 37. Cneoraceae (allied to 36?) 38. Oxalidaceae (allied to 39, 63 and Mimoseae). 39. Geraniaceae (descendants of 27). 40. Linaceae (including Erythroxylon and Humiriaceae ; allied to 38, 39 and 63). Vlll.—MYRTIFLORAE (excluding Onagraceae, Trapa and Halor- rhagidaceae ; descendants of V.) 41. . 1 42. Caryocaraceae (allied to 41, 43 and 54). 43. Rhizophoraceae (including Anisophylleae ; allied to 21, 24, 30, 31, 41—42 and 44). 44. Lythraceae (including Rhynchocalyx, Alzatea, Sonnera- tiaceae ! and Punica ! Descendants of 41?) 45. Myrtaceae (e.xcluding Heteropytis! Descendants of 41). 46. Melastomaceae (descendants of 45). 47. Combretaceae (descendants of 41 ?) 48. Geissolomaceae. 49. Penaeaceae. 50. Ohniaceae. 51. Thymelaeaceae (descendants of 47 ?) 52. Elaeagnaceae (same origin ?) IX.— (near VIII., originated from V.)' ?53. (excluding Parnassia! , Txerba, and Anopterus ? Ribes? Bauera!) 54. (excluding Stylobasium, including Plagio- spermum! Dichotomanthes! and Corynocarpus! Near 26, 34, 35, 41, etc., and originated from V.) ! 55. Anacardiaceae (allied to Pomeae, Amygdaleae and Chry- sobalaneae). 1 56. Sabiaceae (allied to 55). ' It is at present difficult to define this big cohort accurately; it will probably be divided into several after a more exhaustive examination. Phylogenetic System of Flowering Plants. 159 57. Burseraccae (allied to 55 and 56). 58. Simarubaccac (allied to 54—57). 59. Koeberliniaceac (allied to 58 ?) 60. Coriariaceae (allied to 58?) 61. Meliaceae (near 22—24, 62, 72, 117, etc., descended from 21). 62. Rutaceae (including Heteropytis! Near 23, 27, 28, 61, ete., descended from 21). 63. Connaraceae (near 38, 40, 64, etc., descended from 21). 64. Leguminosae (including Morinfja ! Near 54, 61—63. etc., descended from 21). ! 65. Proteaceae (allied to 64 ! and 68 ?) 66. Polygalaceae (allied to 54, 61, 64, 67—70, etc) 67. Trigoniaceae (allied to 66, 68 and 69). 68. Vochysiaceae (allied to 65 ? 66, 67 and 69). 69. Diehapetalaeeae (allied to 65 ?—68). 70. Salvadoraceae (allied to 54, 66—69?) 71. Melianthaceae (allied to 54, 58, 61, 63, 64, 66, 72, etc.) 72. Sapindaceae (including Hippocastaneae; allied to 54, 61, 64, 71, 73, etc.) 73. Aceraceae (allied to 72, 74 and 75 ?) 74. Staphyleaceae (allied to 73, 75 ? and 76). 75. Malpijshiaceae (allied to 73 ? and 76 ?) 76. Celastraceae (allied to 74, 75 ? and 77). 77. Hippocrateaceae (allied to 76). ! 78. Aquifoliaceae (allies or descendants of 80). 79. Cyrillaceae (allied to 78?) 80. Ternstroemiaceae(excluding Bonnetieae ; including Eucry- phia I Medusagyne! Marcgravieae! Pentapbylax ! Tetramerista ? Brexia=Thomassetia, , Roussea and Anopterus ? Stachyurus ? Near VI. and 54, descended from V.) ! 81. Pittosporaceae (allied to 80 and X.) 82. Chlaenaceae (allied to 80 and V.) 83. (allied to 80, 81 and X.) 84. Guttiferae (including Bonnetieae ; allied to 80, 83 and 87). 85. Quiinaceae. 86. Brunelliaceae. 87. Dilleniaceae (allied to V., 80, 83 and 84). X.— (including Primulales, e.xcluding Plumbaginaceae; near 81 and 83, descended from 80). 88. Clethraceae. 89. Pirolaceae. 90. Lennoaceae, 91. Ericaceae (including Vacciniaeeae). ! 92. Empetraceae (descendants, or only a tribe of 91). ! 93. Tremandraceae (including Bauera! Descendants, or only a tribe of 91). 94. Epacridaceae. 95. Diapensiaceae. 96. Theophrastaceae. 97. Myrsinaceae. 98. Primulaceae (connected by Cyclamineae with 97). Hans Hallier.

XI.—SARRACRNIALES ((.Icscendants of X !) 99. Sarraccniaceae (allied to 89, 95, etc.) a. Sarracenieae. h. Ncpentlicae. 100. Droscraccae (including Roridtileae I excltiding Parnassia ! Allied to 89, 93—95, 99, etc.) XII.—SANTALALRS. 101. Olacaceac(incliidinj:;Opilieae! Champcreia! Icacinaceae! Brazzeia ! Erythropyxis ? Rhaptopetaliim ! Egassea Pierre! Scytopetalum! Alanj^itim! Marlea! Bracliy- nema! Ancistrocladtis ! and Sibangea ? Near 22, 23, 2(i, 27, 30, 33, 35, etc., descended froni 21). ! 102. Ampclidaceae (allied to 101, etc.) 103. Grtibhiaceae. 104. Santalaceae. 105. Myzodendraceae. ! 106. Gnetaceae (allied to 104, 105 and 107). 107. Loranthaceae (allied to 101 — 106). X111. — U MBELLIFLORAE. 108. Cornaceae (excludins:; Alangieae! descendants of 101 ?) 109. Araliaceae(incltiding Umbelliferae ; descendants of 108?) 110. Adoraceae (allied to 109?) XIV.—AMENTIFLORAE. ! 111. Hamamelidaceae (including Daphniphylltim and Balanops! Cercidiphyllum, Euptelea atid Eucommia! Platanus! Leitneria ! Buxeae and Stylocereae! Near 26—30, 35, 101, 108, etc. descended frotn V.) 112. Myricaceae (near Leitneria, descended from 111). 113. Salicaceae (near Leitneria, 112 and Daphniphyllum, descended from 111). 114. Juglandaceae (allied to Daphniphyllum, 115a and 116). a. Julianieae. b. Jtiglandeae. 115. Betulaceae (near Corylopsis, Hamamelis, Parrotia, etc., descended from Hamamelidoideae). a. Coryleae. b. Betuleae. c. Casuarineae. 116. Fagaceae (allied to 114 and 115a). XV. PASSIFLORALES (including Campanulatae). 117. Flacourtiaceae (including Lacistema ? near 22—29, descended from 21). 118. Violaceae (allied to 117, 119-121, 125, 131, 132, 138, etc.) 119. Cistaceae (allied to 118). 120. Cochlospermaceae. 121. Bixaceae. 122. Passifloraceae (near 22 and 23, derived from 21). 123. Achariaceae (descendants of 122). 124. JVlalesherbiaceae (same origin). 125. Turneraceae (same origin). 1126. Onagraceae (including Trapa! , near 125, descended from 122). 127. Grossulariaceae (allied to 126?). 128. Elatinaceae (allied to 126 ? ). 129. Datiscaceae (e.xcluding Tetrameleae ?, allied to 126 and 130?). Phylogenetic System of Flowering Plants. i6i ! 130. Halorrbagidaceae (including Hippuris ! and Callitriclie ; descendants of 126 ? ). I 131. Balsaminaceae (near 123, 126, 132, etc., descended from 122). a. Parnassieae; b. Limnantheae (including Mac- gregoria ?); c. Tropaeoleae ; d. Balsamineae. ! 132. Gentianaceae (near 136. 131, 138, etc., derived from 122). ! 133. Aristolochiaceae (near 137, derived from 122). 1 134. Rafflesiaceae (reduced parasitic Aristolocbiaceae). 135. Loasaceae (allied to 126, 136, 137 and 138). 136. Begoniaceae (allied to 135 and 137). 137. Cucurbitaceae (near 133, 135 and 136, originated from 122). 138. (including Stacldiousia? and Peganum! Near 123—126, 131, 132, 135—137, descended from 122). 139. (allied to 138). 140. CandoUeaceae (allied to 138-139). 141. Calyceraceae (allied to 138—140 and 142). 142. Compositae (descendants of 138). XVI.—CENTROSPERMAE. 143. Crassulaceae (including Cepbalotus ; near 132, descended from 122?). I 144. Cactaceae (descendants of 122). I 145. Hydnoraceae (reduced parasitic Cactaceae). ! 146. Balanophoraceae (including Cynomorium ; still more reduced Cactaceae). 147. Aizoaceae (allied to 144). 148. Tbelygonaceae (allied to 147). 149. Portulacaceae (allied to 144, 147—148). 150. Basellaceae (allied to 149). 151. Pbytolaccaceae (allied to 144, 147—150). 1152. Tamaricaceae (including Fouquiera! and Frankeniae! allied to 144). 153. Caryopbyllaceae (allied to 143, 147—152, etc.). 154. Poly^onaceae. ! 155. Plumbaginaceae (allied to 152—154 and 156). 156. Nyctaginaceae (allied to 155). 157. Amarantaceae. 158. Chenopodiaceae. 159. Batidaceae. XVII. CAPRIKOLIALES. 160. Caprifoliaceae (including Sambucus, Silvianthus and Carlemannia,; descendants of 126 ? ). 161. Valerianaceae (descendants of 160). 162. Dipsacaceae (descendants of 161). IVI11. TuBiFLORAE (in a wider sense ; descendants of 21). a. CoNTORTAE (excluding Oleaceac, Salvadoraceae and Gentianaceae). 163. Apocynaceae (including* Asclcpiadaceae ; descended from and by Tabcmacmontana, etc. connected with 21). 164. (including Gelsemium, excluding Budd- leioideae ! Plocosperma ? and Desfontainea I Allied to 163, 165. 166 and 173). 162 Phylogenetic System of Flowering Plants. ! 165. Rubiaceae (excluding Silvianthus 1 and Carlemannia ! Near 163 and t64, descended from 2t ? ). b. TuBiFi.oRAE (in tbe proper sense; excluding Convol- vulaceae !). 166. Bignoniaceae (including Paulownia ! and Wigbtia. Near Hippocastaneae, t63! and 165? descended from 21). ! t67. (reduced Bignoniaceae). 168. Myoporaceae (excluding Zombiana ? andOftia! Reduced Crescentieae ?). 169. Pedaliaceae (including Martyniaceae ; descendants of 166). 170. Acanthaceae (including ZenUerina I Same origin). 171. Verbenaceae (including Phryma ? and Buddleioideae ! Same origin). 172. Labiatae (descendants of 171). 173. Scrophulariaceae (including Plocosperma ? Desfon- tainea! Columellia! Oftia ? Tetracbondra! Globula- rieae I Plantagineae ! Lentibularieae ! Orobancheae !; excluding Paulownia 1 Wightia ! Brookea ! UrosUinnera, Dermatobotrys! Ourisia (partly)! Rebmannia! and Zenkerina, Descendants of 166). 174. Gesneraceae (including Brookea! Uroskinnera, Derma- tobotrys! Ourisia (partly)! and Rehmannia! Close to 173, descended from 166). 175. Solanaceae (excluding Retzia! including Nolaneae! allied to 173), 176. Polemoniaceae (allied 175 and 166). 177. Boraginaceae (excluding Tretracbondra!; close to 171, descended from 166). 178. Hydropbyllaceae (descendents of 177?). /3.—MONOCOTYLEDONS (descendants of 11)'. XIX.—HELOBIAE. 179. Butomaceae (derived from extinct Nympbaeaceae). 180. Alismaceae (allied to 11 and 179). 181. Hydrocbaritaceae (descendants of 180). 182. Potamogetonaceae (allied to 180). 183. Aponogetonaceae (allied to 180 and 182). 184. Juncaginaceae. 185. Triuridaceae. Etc., Etc. ' This division to be worUcd out later. Hamburg, Government Botanical Museum. May 1st, 1905.