ANGIOSPERM CLASSIFICATION
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- Abner Hall
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1 ANGIOSPERM CLASSIFICATION The classification of angiosperms is still very controversial, and two schemes are presented here the first (B&H) as a means to LEARNING the families, the second (APG) as a presentation of the rudiments of an agreed EVOLUTIONARY TREE. Historically a number of schemes have been produced. Bentham & Hooker's Genera Plantarum ( ) comprising 200 families and 7,569 genera pre-dated the publication of Charles Darwin s Origin of Species, so this is essentially a non-evolutionary scheme. However, the arrangement of families was based on characters which provided a sequence from what they took as the most primitive to the most derived families the Scalae Naturae. The classification was based on few, essential characters, and as a result is highly predictive examination of these characters will place an unknown specimen into a particular category. The Angiosperm Phylogeny Group (APG): Systematic research over the past 150 years has changed our view of plant classification, especially in the last 10 years, when major advances in DNA sequencing have provided a completely new avenue of approach. Numerous classifications and schemes have been published: Hutchinson (328 families), Cronquist (388), Takhtajan (592) Dahlgren (477), Thorne (454) Brummitt (445), and Reveal (612), each differ in the recognition of families and the overall scheme. As a result, families, orders and sub-classes are not comparable between any of these systems. Molecular studies are by no means final, and, contrary to what many reference books and journals might imply, a modern synthesis is far from stabilised. However, a consensus is now beginning to emerge. The combined work of many scientists under an umbrella grouping known as the Angiosperm Phylogeny Group (APG) has produced the rudiments of an agreed evolutionary tree. BENTHAM & HOOKER CLASSIFICATION 1. DICOTYLEDONS 2 cotyledons, Exogenous growth POLYPETALAE petals separate 1) THALAMIFLORAE Sepals, Petals and Stamens all attached to receptacle. a) Ranales Gynoecium apocarpous. 1. RANUNCULACEAE; 2. DILLENIACEAE; 3. CALYCANTHACEAE; 4. MAGNOLIACEAE; 5. ANNONACEAE; 6. MENISPERMACEAE; 7. BERBERIDACEAE; 8. NYMPHACEAE. b) Parietales Parietal placentation. [NOT Natural. Convergent evolution: Papaveraceae close to Ranales; remainder scattered amongst Rosids] 9. SARRACENIACEAE; 10. PAPAVERACEAE; 11. CRUCIFERAE; 12. CAPPARACEAE; 13. RESEDACEAE; 14. CISTACEAE; 15. VIOLACEAE; 16. CANELLACEAE; 17. BIXACEAE. c) Polygalineae Ovary 2-3 septate. 18. PITTOSPORACEAE; 19. TREMANDRACEAE; 20. POLYGALACEAE. d) Caryophyllineae Axile placentation. 21. FRANKENIACEAE; 22. CARYOPHYLLACEAE; 23. PORTULACACEAE; 24. TAMARICACEAE. e) Guttiferales Stamens numerous; Calyx imbricate. 25. ELATINACEAE; 26. HYPERICACEAE; 27. GUTTIFERAE; 28. THEACEAE; 29. DIPTEROCARPACEAE; 30. SARCOLAENACEAE. f) Malvales Stamens numerous; Calyx valvate. 31. MALVACEAE; 32. STERCULIACEAE; 33. TILIACEAE. 2) DISCIFLORAE Ovary superior, immersed in disk of flower. a) Geraniales Ovule pendulous, raphe ventral. 34. LINACEAE; 35. HUMIRIACEAE; 36. MALPIGHIACEAE; 37. ZYGOPHYLLACEAE; 38. GERANIACEAE; 39. RUTACEAE; 40. SIMAROUBACEAE; 41. OCHNACEAE; 42. BURSERACEAE; 43. MELIACEAE; 44. DICHAPETALACEAE.
2 b) Olacales Ovule pendulous, raphe dorsal. 45. OLACACEAE; 46. AQUIFOLIACEAE. c) Celastrales Ovule erect, raphe ventral. 47. CELASTRACEAE; 48. STACKHOUSIACEAE; 49. RHAMNACEAE; 50. VITACEAE. d) Sapindales Ovule ascending, raphe ventral to reversed. 51. SAPINDACEAE; 52. MELIOSMACEAE (=Sabiaceae); 53. ANACARDIACEAE; 54. CORIARIACEAE; 55. MORINGACEAE. 3) CALYCIFLORAE Stamens fused to Calyx of flower a) Rosales Ovaries separate, rarely united. [NOT Natural: Saxifragales as Eudicots, Eurosids I, some Asterids] 56. CONNARACEAE; 57. LEGUMINOSAE; 58. ROSACEAE; 59. SAXIFRAGACEAE; 60. CRASSULACEAE; 61. DROSERACEAE; 62. HAMAMELIDACEAE; 63. BRUNIACEAE; 64. HALORAGACEAE. b) Myrtales Ovary syncarpous; divided into locules.[still form a natural group at the base of the Rosids] 65. RHIZOPHORACEAE; 66. COMBRETACEAE; 67. MYRTACEAE; 68. MELASTOMATACEAE; 69. LYTHRACEAE; 70. ONAGRACEAE. c) Passiflorales Ovary syncarpous; Parietal placentation [more or less natural in the Eurosids I] 72. LOASACEAE; 73. TURNERACEAE; 74. PASSIFLORACEAE; 75. CUCURBITACEAE; 76. BEGONIACEAE; 77. DATISCACEAE. d) Ficoidales Ovary syncarous; sub-basal placentation [the basal placentation is critical in placing these families among the Caryophyllids] 78. CACTACEAE; 79. AIZOACEAE. e) Umbellales Ovary syncarpous; 1 ovule per locule. [not wholly natural; these families belong amongst the basal Asterids] 80. UMBELLIFERAE; 81. ARALIACEAE; 82. CORNACEAE GAMOPETALAE petals fused 1) INFERAE Ovary inferior; stamen no. = petal no. a) Rubiales Stamens epipetalous; locules 2-many; ovules 1-many. 83. CAPRIFOLIACEAE; 84. RUBIACEAE. b) Asterales Stamens epipetalous; locule 1; ovule VALERIANACEAE; 86. DIPSACACEAE; 87. CALYCERACEAE; 88. COMPOSITAE. c) Campanulales Stamens free; locules 2-6; ovules many. 89. STYLIDACEAE; 90. GOODENIACEAE; 91. CAMPANULACEAE. 2) HETEROMERAE Ovary superior; stamens opposite petals or double the petal no. a) Ericales Stamens double petal no.; Ovary 2-many locules. 93. ERICACEAE; 93. CLETHRACEAE; 95. EPACRIDACEAE (=Ericaceae); 96. DIAPENSIACEAE; 97. LENNOACEAE (=Boraginaceae). b) Primulales Stamens opposite petals; Ovary 1-locule; Axile placentation. 98. PLUMBAGINACEAE; 99. PRIMULACEAE; 100. MYRSINACEAE. c) Ebenales Stamens opposite petals; Few large seeds SAPOTACEAE; 102. EBENACEAE; 103. STYRACACEAE. 3) BICARPELLATAE Ovary superior, with 2 carpels. a) Gentianales Corolla regular; leaves opposite OLEACEAE; 105. SALVADORACEAE; 106. APOCYNACEAE; 107. ASCLEPIADACEAE; 108. LOGANIACEAE; 109. GENTIANACEAE. b) Polemoniales Corolla regular; leaves alternate POLEMONIACEAE; 111. HYDROPHYLLACEAE; 112. BORAGINACEAE; 113. CONVOLVULACEAE; 114. SOLANACEAE. c) Personiales Corolla irregular to oblique; ovules many.
![3) CALYCIFLORAE Stamens fused to Calyx of flower a) Rosales Ovaries separate, rarely united. [NOT Natural: Saxifragales as Eudicots, Eurosids I, some Asterids] 56. CONNARACEAE; 57. LEGUMINOSAE; 58.](/docs-images/46/21398746/images/page_2.jpg "ROSACEAE; 59. SAXIFRAGACEAE; 60. CRASSULACEAE; 61. DROSERACEAE; 62. HAMAMELIDACEAE; 63. BRUNIACEAE; 64. HALORAGACEAE. b) Myrtales Ovary syncarpous; divided into locules.")
3 115. SCROPHULARIACEAE; 116. GLOBULARIACEAE (=Plantaginaceae); 117. LENTIBULARIACEAE; 119. GESNERIACEAE; 120. BIGNONIACEAE; 121. PEDALIACEAE; 122. ACANTHACEAE. d) Lamiales Corolla irregular to oblique; 1-2 ovules MYOPORACEAE; 125. VERBENACEAE; 126. LABIATAE; 127. PLANTAGINACEAE MONOCHLAMYDEAE only 1 kind of perianth (not petals + sepals) This division was never intended to be natural it was merely to help identify these singleperianthed plants. Some are basal families (Aristolochia, Piper) with unusual flower structures, whilst others are highly derived, with sepals or petals lost during evolution. Biochemistry solved the question of the placement of many of these families. 1) Curved embryos [The curved embryos & seeds make these part of the Caryophyllids] 128. NYCTAGINACEAE; 130. AMARANTHACEAE; 131. CHENOPODIACEAE; 133. BATIDACEAE; 134. POLYGONACEAE. 2) Many-seeded aquatics ( Caryophyllids / Palaeoherbs) 135. PODOSTEMACEAE; 136. NEPENTHACEAE; 137. RAFFLESIACEAE. 3) Microscopic seeds ( Paleotrees/Palaeoherbs) 138. ARISTOLOCHIACEAE; 139. PIPERACEAE; 140. CHLORANTHACEAE; 141. MYRISTICACEAE; 142. MONIMIACEAE. 4) Daphnales ( Paleotrees/Eudicots/Rosids) 143. LAURACEAE; 143. HERNANDIACEAE; 144. PROTEACEAE; 145. THYMELAEACEAE; 146. PENAEACEAE; 147. ELAEGNACEAE. 5) Achlamydosporaceae ( Eudicots/Asterids) 148. LORANTHACEAE; 149. SANTALACEAE; 150. BALANOPHORACEAE. 6) Unisexulaes ( Eudicots/Rosids) 151. EUPHORBIACEAE; 152. BALANOPACEAE; 153. URTICACEAE; 154. PLATANACEAE; 155. LEITNERIACEAE; 156. JUGLANDACEAE; 157. MYRICACEAE; 158. CASUARINACEAE; 159. BETULACEAE. 7) Ordines anomalus ( Paleoherbs/Rosids) 160. SALICACEAE; 162. EMPETRACEAE; 163. CERATOPHYLLACEAE. 2. MONOCOTYLEDONS 1 cotyledon, Endogenous growth. 1) Microscopic seeds 167. HYDROCHARITACEAE; 168. BURMANNIACEAE; 169. ORCHIDACEAE. 2) Inferior ovaries 170. ZINGIBERACEAE; 171. BROMELIACEAE; 172. HAEMODORACEAE; 173. IRIDACEAE; 174. AMARYLLIDACEAE; 175. VELLOZIACEAE; 176. DIOSCOREACEAE. 3) Corolla-like flowers 177. STEMONACEAE; 178. LILIACEAE; 179. PONTEDERIACEAE; 180. PHILYDRACEAE; 181. XYRIDACEAE; 182. MAYACACEAE; 183. COMMELINACEAE; 184. RAPATEACEAE. 4) Calyx-like flowers 185. FLAGELLARIACEAE; 186. JUNCACEAE; 187. PALMAE. 5) Perianth-less flowers 188. PANDANACEAE; 189. CYCLANTHACEAE; 190. TYPHACEAE; 191. ARACEAE; 192. LEMNACEAE. 6) Apocarpous 193. TRIURIDACEAE; 194. ALISMATACEAE; 195. JUNCAGINACEAE. 7) Glumaceous flowers 196. ERIOCAULACEAE; 197. CENTROLEPIDACEAE; 198. RESTIONACEAE; 199. CYPERACEAE; 200. GRAMINEAE.
4 ANGIOSPERM PHYLOGENY GROUP AN APPROXIMATE EVOLUTIONARY TREE The branching lines demonstrate how closely related different families are. Each group of families are arranged alphabetically, those families in bold are grown at Glasnevin, those underlined represent order names (which end in ales). Arrows indicate continuations to other pages. One important revelation of the new synthesis is that there are not just Monocots and Dicots as two classes of Angiosperms, rather the earliest evolution of flowering plants led to four basic lineages: PALEOHERBS, represented by Waterlilies, Aristolochiaceae, Piperaceae, Schisandra. PALEOTREES, represented by Magnoliaceae, Winteraceae, Lauraceae. MONOCOTS, the traditional Monocots. EUDICOTS, so called because they are proper or true dicots without the paleo-s above. A pattern of groupings is discernible within the Eudicots: The most ancient grouping in this lineage are indeed the Ranunculaceae, which have always been traditionally viewed as the progenitors of all other flowering plants, then a number of distinct basal groups such as the Caryophyllids and Saxifragales can be distinguished, followed by two very distinctive groups, the Rosids and Asterids. Within these latter two groups we can recognise a similar pattern of evolutionary left-overs basal groups and distinguished more clearly from these, clusters of closely-related families, which are termed Eu-Rosids and Eu-Asterids respectively. Amborellaceae Cabombaceae Chloranthaceae Nymphaeaceae Austrobaileyaceae Illiciaceae Schisandraceae Ceratophyllaceae PALEOHERBS Aristolochiaceae Lactoridaceae Piperaceae Saururaceae Cannelaceae Winteraceae Atherospermataceae Calycanthaceae Gomortegaceae Hernandiaceae Lauraceae Monimiaceae Annonaceae Degeneriaceae Eupomatiaceae Himantandraceae Magnoliaceae Myristicaceae PALEOTREES EUDICOTS MONOCOTS
5 EUDICOTS (Tricolpate Pollen) Berberidaceae Circaeasteraceae Eupteleaceae Lardizabalaceae Menispermaceae Papaveraceae [+Fumariaceae] Ranunculaceae Nelumbonaceae Platanaceae Proteaceae Achatocarpaceae Aizoaceae Amaranthaceae Asteropeiaceae Basellaceae Cactaceae Caryophyllaceae Chenopodiaceae Didiereaceae Molluginaceae Nyctaginaceae Buxaceae Frankeniaceae Physenaceae Sabiaceae Tamaricaceae Phytolaccaceae Portulacaceae Rhabdodendraceae Olacaceae Opiliaceae Loranthaceae Misodendraceae Santalaceae Altingiaceae Cercidiphyllaceae Crassulaceae Daphniphyllaceae Grossulariaceae Haloragaceae Hamamelidaceae Iteaceae Paeoniaceae Penthoraceae Pterostemonaceae Saxifragaceae Tetracarpaeaceae SAXIFRAGALES Sarcobataceae Simmondsiaceae Stegnospermataceae Ancistrocladaceae Dioncophyllaceae Droseraceae Drosophyllaceae Nepenthaceae Plumbaginaceae Polygonaceae Curved Embryos Straight Embryos CARYOPHYLLIDS ROSIDS ASTERIDS Aextoxicaceae Berberidopsidaceae Dilleniaceae Gunneraceae Myrothamnaceae Vitaceae Note: Underlined family names represent Order names for the groups. i.e. Caryophyllales, Saxifragales etc.
6 ROSIDS Francoaceae Caryocaraceae Geraniaceae Chrysobalanaceae Greyiaceae Clusiaceae Ledocarpaceae Dichapetalaceae Melianthaceae Erythroxylaceae Celastraceae Euphorbiaceae Parnassiaceae Euphroniaceae Stackhousiaceae Flacourtiaceae Alzateaceae Hugoniaceae Combretaceae Irvingiaceae Crypteroniaceae Linaceae Heteropyxidaceae Malpighiaceae Lythraceae Ochnaceae Melastomataceae Pandaceae Myrtaceae Passifloraceae Oliniaceae Quiinaceae Onagraceae Rhizophoraceae Penaeaceae Salicaceae Psiloxylaceae Turneraceae Vochysiaceae Violaceae Akaniaceae Cephalotaceae Bataceae Connaraceae Brassicaceae [Capparaceae] Cunoniaceae Caricaceae Elaeocarpaceae Emblingiaceae Oxalidaceae Gyrostemonaceae Koeberliniaceae Fabaceae Limnanthaceae Polygalaceae Moringaceae Quillajaceae Pentadiplandraceae Surianaceae Resedaceae Salvadoraceae Barbeyaceae Setchellanthaceae Cannabaceae Tovariaceae Cecropiaceae Tropaeolaceae Celtidaceae Elaeagnaceae Bixaceae Moraceae Cistaceae Rhamnaceae Cochlospermaceae Rosaceae Dipterocarpaceae Ulmaceae Malvaceae Urticaceae Muntingiaceae Sarcolaenaceae Begoniaceae Sphaerosepalaceae Coriariaceae Sterculiaceae Cucurbitaceae Thymelaeaceae Datiscaceae Tiliaceae Tetramelaceae Anacardiaceae Betulaceae Biebersteiniaceae Casuarinaceae Burseraceae Fagaceae Kirkiaceae Juglandaceae Meliaceae Myricaceae Rutaceae Nothofagaceae Sapindaceae [Aceraceae] Simaroubaceae EUROSIDS I EUROSIDS II
7 Cornaceae [+Nyssaceae ] Grubbiaceae Hydrangeaceae Hydrostachyaceae Loasaceae ASTERIDS Actinidiaceae Balsaminaceae Clethraceae Diapensiaceae Ebenaceae Ericaceae Lecythidaceae Marcgraviaceae Myrsinaceae Polemoniaceae Primulaceae Roridulaceae Sapotaceae Sarraceniaceae Styracaceae Symplocaceae Ternstroemiaceae Theaceae Theophrastaceae SUPERIOR OVARY ANTHER no. = or greater than Petal no. Boraginaceae Hydrophyllaceae Plocospermataceae Vahliaceae Aucubaceae Eucommiaceae Garryaceae Oncothecaceae Apocynaceae Gelsemiaceae Gentianaceae Loganiaceae Rubiaceae Aquifoliaceae Helwingiaceae Acanthaceae Phyllonomaceae Avicenniaceae Bignoniaceae Apiaceae Buddlejaceae Araliaceae Byblidaceae Aralidiaceae Gesneriaceae Griseliniaceae Lamiaceae Melanophyllaceae Lentibulariaceae Pittosporaceae Myoporaceae Torricelliaceae Oleaceae Orobanchaceae Caprifoliaceae Paulowniaceae Diervillaceae Pedaliaceae Dipsacaceae Phrymaceae Linnaeaceae Plantaginaceae Morinaceae Schlegeliaceae Valerianaceae Scrophulariaceae Stilbaceae Alseuosmiaceae Tetrachondraceae Argophyllaceae Verbenaceae Bruniaceae Asteraceae Escalloniaceae Calyceraceae Convolvulaceae Icacinaceae Campanulaceae Hydroleaceae [+Lobeliaceae] Montiniaceae Goodeniaceae Solanaceae Menyanthaceae Sphenocleaceae Pentaphragmataceae Stylidiaceae EUASTERIDS II : INFERIOR OVARY EUASTERIDS I : SUPERIOR OVARY ANTHER no. = or fewer than Petal no.
8 MONOCOTS Acoraceae Orchidaceae Araceae Iridaceae Alismataceae Aponogetonaceae Hemerocallidaceae Butomaceae Phormiaceae Hydrocharitaceae Juncaginaceae Hyacinthaceae Limnocharitaceae Themidaceae Posidoniaceae Potamogetonaceae Agapanthaceae Ruppiaceae Boryaceae Alliaceae Scheuchzeriaceae Amaryllidaceae Tofieldiaceae Zosteraceae Agavaceae Anthericaceae ASPARAGALES Asparagaceae Asphodelaceae Asteliaceae Convallariaceae Doryanthaceae Ruscaceae Xanthorrhoeaceae Burmanniaceae Dioscoreaceae Taccaceae Cyclanthaceae Pandanaceae Stemonaceae Velloziaceae Campynemataceae Liliaceae Philesiaceae Ripogonaceae Smilacaceae LILIALES Alstroemeriaceae Colchicaceae Melanthiaceae (Trilliaceae) Arecaceae Bromeliaceae COROLLA & CALYX NOT ALIKE Stems WITH VESSELS Anarthriaceae Centrolepidaceae Cyperaceae Eriocaulaceae Flagellariaceae Joinvilleaceae Juncaceae Poaceae Restionaceae Sparganiaceae Typhaceae Commelinaceae Haemodoraceae Philydraceae Pontederiaceae Cannaceae Costaceae Heliconiaceae Lowiaceae Marantaceae Musaceae Strelitziaceae Zingiberaceae
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