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THE

VOL. 58

BOTANICAL
JULY-SEPTEMBER, 1992

REVIEW
NO. 3

Classification and Geography of the Flowering Plants


ROBERT F. THORNE
Rancho Santa Ana Botanic Garden Claremont, California 91711

Dedication
This long review paper on the classification and geography of the flowering plants is dedicated to the memory of Arthur Cronquist, the late editor of Botanical Review, who requested this paper and encouraged its production. Arthur was a good friend, fine field man, highly intelligent scholar, prodigious worker, and productive author. Although we sometimes did not agree on matters of angiosperm phylogeny, we greatly enjoyed our verbal jousting and remained close friends. Arthur's recent death is a shocking loss to the fields of floristics, plant phylogeny, and phytogeography. He will be greatly missed by his colleagues, friends, and other admirers in the botanical world. I. Abstract .................................................................................................................................................................................. 226 Resumen ............................................................................................................................................................................... 227 II. Introduction ........................................................................................................................................................................ 227 A. Important Literature Subsequent to 1981 .......................................................................................... 227 1. Inclusive Works for Phylogeny ....................................................................................................... 227 2. Floral Morphology .................................................................................................................................... 228 3. Palynology ....................................................................................................................................................... 229 4. Seed and Fruit Morphology ............................................................................................................... 230 5. Embryology .................................................................................................................................................... 230 6. Stem Anatomy ............................................................................................................................................. 231 7. Leaf Architecture ....................................................................................................................................... 232 8. Karyomorphology ..................................................................................................................................... 232 9. Micromorphology ...................................................................................................................................... 232 10. Chemotaxonomy and Molecular Taxonomy ........................................................................ 233 11. Phytoserology ................................................................................................................................................ 234 12. Paleobotany .................................................................................................................................................... 235 a. Origins and Early Evolution of Angiosperms ............................................................... 235 b. Tertiary Paleobotany ....................................................................................................................... 237 13. Phytogeography ........................................................................................................................................... 238

Copies of this issue [58(3)] may be purchased from the Scientific Publications Department, The New York Botanical Garden, Bronx, NY 10458-5126 USA. Please inquire as to prices.

The Botanical Review 58: 225-348, Jul.-Sept., 1992 9 1992 The New York Botanical Garden

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THE BOTANICAL REVIEW 14. Host-Parasite Relationships .............................................................................................................. 240 15. Cladistics 240 Classification of the Angiospermae ................................................................................................................... 241 A. Changes Made in the System Since 1983 ........................................................................................... 241 I. Dicotyledoneae ............................................................................................................................................ 241 2. Monocotyledoneae .................................................................................................................................... 243 B. Explanation of the Phyletic Shrub ........................................................................................................... 244 C. Explanation of the Synopsis .......................................................................................................................... 244 1. Philosophy of Classification .............................................................................................................. 244 2. Nomenclature ............................................................................................................................................... 245 3. Recent Monographs and Revisions ............................................................................................. 245 4. Symbols and Numbers .......................................................................................................................... 246 5. Distribution of Taxa ................................................................................................................................ 247 6. Arrangement of Taxa in Synopsis ................................................................................................. 248 7. Discussion of Tables ............................................................................................................................... 248 Classification and Distribution of the Families and Subfamilies of the Class Angiospermae (Magnoliopsida) ......................................................................................................................................... 258 Acknowledgments .......................................................................................................................................................... 293 Literature Cited ................................................................................................................................................................ 294 Index to Scientific Names ........................................................................................................................................ 327 Index to Common Names ....................................................................................................................................... 347
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III.

IV. V. VI. VII. VIII.

I. Abstract
Thorne, Robert F. (Rancho Santa Ana Botanic Garden, Claremont, CA 91711). Classification and geography of the flowering plants. Bot. Rev. 58(3): 225-348. 1992.This treatment of the flowering plants is the latest revision of my classification of the Class Angiospermae and replaces my 1983 and more recent 1992 synopses. An update is necessary because so much new information has been published in the last decade pertinent to the classification of the flowering plants. About 870 such recent books, monographs, and other botanical papers are cited in the Introduction, listed primarily by the botanical discipline that they represent. Also considerable changes in my classification have been necessitated by my narrowed family- and ordinal-gap concepts, acceptance of the ending "-anae" for superorders in place of the traditional but inappropriate "-iflorae," and acceptance of more prior or more widely used names for the categories above the family. A new phyletic "shrub" replaces earlier versions, and attempts to indicate visually relative sizes and relationships among the superorders, orders, and suborders. One table includes a statistical summary of floweringplant taxa: ca. 233,900 species of 12,650 genera, 437 families, and 708 subfamilies and undivided families in 28 superorders, 71 orders, and 71 suborders of Angiospermae. Three other tables summarize the known indigenous distribution of the families and subfamilies of angiosperms about the world. The synopsis lists the flowering plant taxa from the class down to the subfamily (and in Asteraceae down to the tribe) with indication of the degree of confidence I place in the circumscription and placement of each category above the subfamily, the best available estimates of the number of genera and species for each category, and the known indigenous distribution of each subfamily and family. Table V lists alphabetically the geographical abbreviations used in the synopsis. The extensive bibliography of recent literature should be helpful to those persons interested in the classification of the flowering plants.

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Resumen
Este tratado de las plantas con flores es la revisi6n mils reciente de mi clasificaci6n de la Clase Angiospermae y substituye mis sinopsis de 1983 y la reciente de 1992. Este modificacion es necesaria debido a la gran cantidad de informaci6n nueva que se ha publicado en la 61tima decada, acerca de la clasificaci6n de las plantas con flores. En la introducci6n de esta revisirn se citan alrededor de 870 trabajos recientes los cuales incluyen libros, monografias y otros articulos, que son listados de acuerdo con la disciplina que representan. Asimismo, ciertos cambios significativos en mi clasificaci6n son el resultado de mi concepto estricto a mivel familia y mis conceptos de separaci6n entre ordenes, la aceptaci6n de la terminaci6n "-anae" para los super6rdenes, en lugar de la tradicional pero inapropiada "-iflorae"; y la aceptaci6n, ya sea de hombres anteriores o hombres m~is comfinmente aceptados para las categorias a nivel familia o a nivel superior. E1 nuevo arbusto filrtico substituye las versiones recientes e intenta indiear visualmente los tamafios proporcionales y las relaciones entre super6rdenes, 6rdenes y sub6rdenes. Se presenta una tabla con un sumario de las estadisticas de los taxa comprendidos dentro de las plantas con flores: alrededor de 233,900 especies en 12,650 grneros, 437 familias, 708 subfamilias y familias no subdivididas, todos 6stos incluidos en 28 super6rdenes, 71 6rdenes y 71 sub6rdenes de Angiospermae. En otras tres tablas que se presentan en este trabajo se sumariza la distribuci6n indigena de las familias y subfamilias de angiospermas en el mundo. Esta sinopsis lista los taxa incluidos en las plantas con flores desde el nivel orden hasta el nivel de subfamilia (en Asteraceae hasta tribu), indicando el grado de confianza que designo a la circunscripci6n y a la localizaci6n de cada categoria al nivel de subfamilia o superior. Tambirn incluyo las estimaciones disponibles acerca del nfimero de grneros y especies para cada categoria y la distribuci6n geogr~fica indigena conocida de cada subfamilia y familia. La tabla 5 lista alfabrticamente las abreviaciones geogr~ficas usadas en esta sinopsis. La extensa bibliografia de la literatura reciente incluida en este trabajo puede ser de utilidad para todo interesado en la clasificaci6n de las plantas con flores.

II. Introduction Nearly a decade has passed since I published in the Nordic Journal of Botany my "Proposed new realignments in the angiosperms" (Thorne, 1983). In that time hundreds of papers and many books have been published on the paleontology, morphology, palynology, phytochemistry, karyomorphology, molecular taxonomy, etc. of the flowering plants.
A. IMPORTANT LITERATURE SUBSEQUENT TO 1981

I. Inclusive Works for ehylogeny


Among the more significant longer publications for the better understanding of angiosperm phylogeny are two volumes on the Anatomy of the Dicotyledons, Second Edition (Metcalfe, 1987; Metcalfe & Chalk, 1983); Chemistry and Chemical Tax-

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onomy of the Rutales, edited by Waterman and Grundon (1983); Proteins andNucleic Acids in Plant Systematics, edited by Jensen and Fairbrothers (1983); The Biology of Mistletoes, edited by Calder and Bernhardt (1983); the Annals of the Missouri Botanical Garden symposium number on the Myrtales (in 1984); Embryology of Angiosperms, by Johri (1984); the monumental work on monocot families by Dahlgren, Clifford, and Yeo (1985); The Chemistry and Biology of lsoquinoline Alkaloids, edited by Phillipson et al. (1985); two important volumes, Floristic Regions of the World and Systema Magnoliophytorum, by Takhtajan (1986, 1987); Genera Graminum: Grasses of the World, by Clayton and Renvoize (1986); Genera Cyperacearum, by Goetghebeur (1986); The Botany of Mangroves, by Tomlinson (1986); Genera Palmarum, by Uhl and Dransfield (1987); monographs on the fossil history of the Juglandaceae by Manchester (1987b) and of Acer by Wolfe and Tanai (1987); The Plant-book, by Mabberley (1987); The Origin ofAngiosperms and their Biological Consequences, edited by Friis et al. (1987); Grass Systematics and Evolution, edited by Soderstrom et al. (1987); the volume on angiosperm evolution and classification by Cronquist (1988); the discussion of families endemic to South Africa by Dahlgren and van Wyk (1988); the review of the Cornaceae and associated families by Eyde (1988); The Flavonoids, edited by Harborne (1988); Advances in Legume Biology, edited by Stirton and Zarucchi (1989); the Reading Symposium volume on the Hamamelididae edited by Crane and Blackmore (1989); two papers on the Dahlgren system by Gertrud Dahlgren (1989a, 1989b); a review of the paleobiogeographic relationships of angiosperms from the Cretaceous and Early Tertiary of the North American area by Taylor (1990); Biology and Utilization of the Cucurbitaceae, edited by Bates et al. (1990); the Annals of the Missouri Botanical Garden symposium number on "Systematics and Evolution of the Monocotyledons" (1990); and the series of Aliso papers (1989-1991) originally presented in the Rolf Dahlgren Memorial Symposium at Berlin in 1987. 2. Floral Morphology
Numerous shorter studies, revisions, and monographs have also expanded very considerably our knowledge of the paleontologic history, evolution, and classification of the angiosperms. Floral anatomy and morphology still remain very useful in the circumscription of taxa and the search for relationships, especially when combined with consideration of embryology, palynology, and seed and fruit morphology. Among the more phylogenetically informative recent morphological studies, listed by the taxa investigated, are those for Acoraceae (Grayum, 1987a), Amaranthaceae (Eliasson, 1988; Volgin, 1987), Anacardiaceae (Wannan & Quinn, 1991), Anisophylleaceae and Rhizophoraceae (Dahlgren, 1988; Tobe & Raven, 1988a, 1988b), Apocynaceae (Fallen, 1986, 1987), Apostasioideae of Orchidaceae (Kumar & Manilal, 1988), Araceae (French, 1986), Asteraceae (Stuessey & Spooner, 1988), Nandina and Berberidaceae (Terabayashi, 1983, 1985), Bixa and Cochlospermum (Decraene, 1989), Boraginaceae (A1-Nowaihi et al., 1987), Bromeliaceae (Brown & Gilmartin, 1989), Brunoniaceae (Erbar & Leins, 1988), Campynemataceae (Goldblatt, 1986b), Caryocaraceae (Dickison, 1990c), Caryophyllales (Brown & Varadarajan, 1985), Casuarinaceae (Dilcher et al., 1990; Flores & Moseley, 1990), Chloranthaceae (Endress, 1987), Circaeasteraceae (Hu, 1987; Hu & Tian, 1985; Hu et al., 1990), Dipsacales (Wagenitz & Laing, 1984), Eriocaulaceae (Stuetzel, 1985), Fabaceae (Tucker, 1989), Fagaceae (Kaul, 1986),

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Greyiaceae (Steyn et al., 1987), Hamamelidaceae (Bogle, 1986, 1989; Endress, 1989a, 1989b, 1989c), Hamamelididae (Dickison, 1989a; Hufford & Endress, 1989), Hydrastidaceae (Tobe & Keating, 1985), Kingdonia of the Anemoneae (Kosuge et al., 1989), Lacandonia (Marquez-Guzman et al., 1989), Lardizabalaceae (Qin, 1989), Lilaea (Posluszny et al., 1986), Magnoliidae (Endress, 1990; Endress & Hufford, 1989), Malpighiaceae (Vogel, 1987), Medusagynaceae (Dickison, 1990a, 1990b), Melanthiaceae (Ambrose, 1985), Menyanthaceae (Ornduff, 1988), monocot septal nectaries (Schmid, 1985), Myrtales (Decraene & Smets, 199 l a), Nelumbonaceae (Ito, 1986; Moseley & Uhl, 1985), Nesogenaceae (Marais, 1981), Nyctanthes in Oleaceae (Kiew & Baas, 1984), Oxalidaceae (Lack & Kevan, 1987), Petiveriaceae (Volgin, 1988), Picrodendron of Euphorbiaceae (Hakki, 1985), Lithophytum of Plocospermeae, Loganiaceae (Chiang & Frame, 1987), Potamogetonaceae (Guo & Cook, 1990), Restionaceae (Kircher, 1986), 'Lower' Rosidae (Endress & Stumpf, 1991), Rutaceae and Simaroubaceae (Ramp, 1988), Salicaceae (Hong &Chen, 1987), Saururaceae (Liang & Tucker, 1989), Schlegelieae in Scrophulariaceae (Armstrong, 1985), Staphyleaceae (Dickison, 1986a; Ramp, 1987), Theophrastaceae (Stahl, 1987), Ticodendron (Tobe, 1991), Tovaria of Capparaceae (Fisel & Weberling, 1990), Trochodendraceae (Endress, 1986), Triuridaceae (Rfibsamen-Weustenfeld, 1991), Zingiberales (Kirchoff, 1988), and Zygophyllaceae (Decraene & Smets, 1991b). 3. Palynology Especially popular in recent years have been palynological investigations. Nowicke, Skvarla, and their associates have been particularly prolific in their production of informative pollen studies. Groups most recently examined by them include the Achatocarpaceae (Skvarla & Nowicke, 1982), Actinidiaceae and Dilleniaceae (Dickison et al., 1982), Boraginaceae (Nowicke & Miller, 1987), Circaeasteraceae and Sargentodoxa (Nowicke & Skvarla, 1982), Corynocarpaceae (Nowicke & Skvarla, 1983b), Gonystylaceae (Nowicke et al., 1985), Helleborus (Nowicke & Skvarla, 1983a), Hoplestigmataceae (Nowicke & Miller, 1989), Loasaceae (Poston & Nowicke, 1990), Lythraceae (Graham et al., 1990), Myrtales (Patel et al., 1984), Paeonia (Nowicke et al., 1986), Panda (Nowicke, 1984), and Simrnondsia (Nowicke & Skvarla, 1984). Other taxa whose circumscription and/or phyletic position have been clarified by palynological studies include the Actinidiaceae (Zhang, 1987), Agavaceae (Alvarez & Koehler, 1987), Alismatales (Chanda et al., 1988), Annonaceae (Hesse et al., 1985; Waha & Morawetz, 1988), Anthericaceae (Roth et al., 1987), Araceae (Grayum, 1987b), Bombacaceae (Nilsson & Robyns, 1986), Cabombaceae (Osborn et al., 1991), Carpodetus (Praglowski & Grafstrom, 1985), Chloanthaceae (Raj & E1-Ghazaly, 1987; Raj & Grafstrom, 1984), Chloranthaceae (Chapman, 1987), Cneoraceae (LobreauCallen & Jeremie, 1986), Corynocarpaceae (Philipson, 1987a), Ctenolophonaceae (Ham, 1989), Desfontainiaceae (Hoc & Bravo, 1984), Diapensiaceae (Xi & Tang, 1990), Empetraceae (Kim et al., 1988), Ericaceae (Takahashi, 1988), Euphorbiaceae (Punt, 1987), Eupomatiaceae (Woodland & Garlick, 1982), Fabaceae (Guinet & Ferguson, 1989), Gunneraceae (Jarzen & Dettmann, 1989), Haemodoraceae and Lanaria, Philydraceae, Tecophilaeaceae, and Pontederiaceae (Simpson, 1983, 1985a, 1985b, 1987), Illiciaceae (Liu & Yang, 1989), Ixioideae of Iridaceae (Goldblatt et al., 1991), Lactoridaceae (Zavada & Taylor, 1986), Lamiaceae (Abu-Asab & Cantino, 1987, 1989; Chadwell et al., 1992), Lennoaceae (Yatskeivych & Zavada, 1984),

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Lythraceae and Alzatea (Graham et al., 1985), Malpighiaceae (Yunus, 1990), Malvaceae (Christensen, 1986), Martyniaceae vs. Pedaliaceae (Bretting & Nilsson, 1988), Melanthiaceae (Takahashi & Kawano, 1989), Monimiaceae and Atherospermatoideae (Lorence et al., 1984; Sampson & Foreman, 1988), Monocotyledoneae (Zavada, 1983b), Monotropoideae ofEricaceae (Takahashi, 1987), Moreae of Moraceae (Punt & Eetnerink, 1982), Morinaceae (Verlaque, 1983), Moringaceae (Ferguson, 1985), Nesogenaceae (Raj, 1985), Oncothecaceae (Dickison, 1986b), Opiliaceae (Hiepko & Lobreau-Callen, 1987), Orchidaceae (Freudenstein, 1988), Orobanchoideae of Scrophulariaceae (Minkin & Eshbaugh, 1989), Pandanaceae (Cox, 1990), Potamogetonaceae (Sorsa, 1988), Proteaceae (Feuer, 1986, 1990), Ratttesiaceae (Takhtajan et al., 1985), Restionaceae (Linder, 1987; Linder & Ferguson, 1985), Sapotaceae (Harley, 1986, 1991), Staphyleaceae (Dickison, 1987a; Long et al., 1985), Stilbaceae (Raj, 1983), Styracaceae (Li & Yu, 1985), Symphoremaceae (Raj, 1984), Swartzioideae of Fabaceae (Ferguson & Skvarla, 1988; Tucker, 1988), Ticodendraceae (Feuer, 1991), Tribuloideae of Zygophyllaceae (Praglowski, 1987), Trimeniaceae and Calycanthaceae (Sampson, 1987; Sampson & Endress, 1984), Ulmaceae (Takahashi, 1989; Zavada, 1983a), Verbenaceae (Raj, 1987), Winteraceae (Doyle et al., 1990a, 1990b), and Zingiberales (Kress, 1987).

4. Seed and Fruit Morphology


Investigations of seed and fruit morphology are also really extensions of floral morphology or possibly even of embryology depending upon one's primary interests. Regardless of labelling, the following taxa have been clarified through such studies: Amaryllidaceae (Oganezova, 1990), Anacardiaceae (Wannan & Quinn, 1990), Aristolochiaceae (Huber, 1985), Asphodelaceae (Oganezova, 1987), Begoniaceae (Bouman & Lange, 1983), Pitcairnioideae of Bromeliaceae (Varadarajan & Gilmartin, 1988), Buddlejaceae (Norman, 1987), Campanulaceae (Shetler & Morin, 1986), Ceratophyllaceae (Les, 1986, 1988a, 1988b, 1989; Wilmot-Dear, 1985), Cneoraceae (Boesewinkel, 1984), Cartonematoideae of Commelinaceae (Grootjen, 1983), Cunoniaceae (Dickison, 1984), Droseraceae (Boesewinkel, 1989), Gesneriaceae (Beaufort-Murphy, 1983), Hernandiaceae (Mohana Rao, 1986), Humiriaceae and Hypseocharis(Boesewinkel, 1985, 1988), Hydrocharitaceae, including Najas (Shaffer-Fehre, 1991a, 1991b), Iridaceae (Manning & Goldblatt, 1991), Juglandaceae (Schaarschmidt, 1985), Loasaceae (Hufford, 1988), Pteridophylloideae of Papaveraceae (Brueckner, 1985), Poaceae (Semikhov, 1988), Polygalaceae (Verkerke, 1984, 1985, 1991), Portulacaceae (Nyananyo, 1988), Potamogetonaceae (Takaso & Bouman, 1984), Rattlesiaceae (Bouman & Meijer, 1987), Rapateaceae (Venturelli & Bouman, 1988), Maloideae of Rosaceae (Rohrer et al., 1991), Solanales (Cocucci, 1983), Sphenocleaceae (Haridasan & Mukherjee, 1988), TovariaofCapparaceae (Boesewinkel, 1990), Trigoniaceae and Vochysiaceae (Boesewinkel, 1987; Boesewinkel & Venturelli, 1987), Ulmaceae (Tobe, 1987), and evolutionary trends in dicotyledonous seeds (Teichman & van Wyk, 1991).

5. Embryology
Embryological investigations, in addition to those listed above that are restricted more specifically to pollen grains, seeds, and fruits, continue to be most helpful to the phylogenists. Some of those that I found especially elucidative are the following listed taxa: Acoraceae and Araceae (Grayum, 1991), Anisophylleaceae (Tobe & Ra-

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ven, 1988a, 1988b), Betulaceae (Chen et al., 1990), Bretschneidera (Tobe, 1990), Brexia (Kamelina, 1988), Burmanniaceae and Corsiaceae (Riibsamen, 1986), Campynemataceae (Dahlgren & Lu, 1985), Chrysobalanaceae and Crypteroniaceae (Tobe & Raven, 1984a, 1987a), Cypripedioideae of Orchidaceae (Sood & Mohana Rao, 1988), Dicotyledoneae (Dahlgren, 1991), Ecdeiocoleaceae (Rudall, 1990), Eriospermaceae (Lu, 1985), Flagellariaceae and Restionaceae (Rudall & Linder, 1988), Gyrostemonaceae (Tobe & Raven, 1991), Haemodoraceae (Steinecke & Hamann, 1989), Lachnanthes of Haemodoraceae (Simpson, 1988), Heteropyxis and Psiloxylon of Myrtaceae (Tobe & Raven, 1987b, 1990), Himantandraceae (Prakash et al., 1984), Loganiaceae (Andronova, 1988), Nandinoideae of Berberidaceae (Ehdaie & Russell, 1985), Petermanniaceae (Conran, 1988), Poaceae (Bhanwra, 1988; Bhanwra et al., 1991), Rapateaceae and Xyridaceae (Tiemann, 1985), Rhabdodendraceae (Tobe & Raven, 1989), Stegnospermataceae (Narayana & Narayana, 1986; Satyanarayana & Narayana, 1985), and Retzia of Stilbaceae (EngeU, 1987). Nagendran and Dinesh (1989) have produced a most useful compendium of embryological titles listed by angiosperm families. 6. Stem Anatomy Another rich source of comparative information for phylogenists continues to be stem anatomical studies. The most prolific author of such informative anatomical publications is my colleague Sherwin Carlquist with the following taxa investigated by him and his associates: Acanthaceae (Carlquist & Zona, 1988a), Asteridae (1990b, 1991 b), Begoniaceae (1985a), Bruniaceae (1991 b), Buxaceae (1982), Chloranthaceae (1987e, 1990d), Cneoraceae (1988a), Convolvulaceae (Carlquist & Hanson, 1991), Coriariaceae (1985 b), Degeneriaceae (1989c), Duckeodendraceae and Goetzeaceae (1988c), Elatinaceae (1984b), Empetraceae (1989d), Gentianaceae (1984a), Geraniaceae (1985d), Gesneriaceae (Carlquist & Hoekman, 1986a), Goetzeaceae (Zona, 1989), Hydrophyllaceae (Carlquist & Eckhart, 1984), Lactoridaceae (1990a), Lamiaceae (1992a), Loasaceae (1984e, 1987d), Malesherbiaceae (1984d), Martyniaceae and Pedaliaceae (1987c), Misodendraceae (1985e), Montiniaceae (1989b), Myoporaceae (Carlquist & Hoekman, 1986b), Nolanaceae (1987a), Papaveraceae (Carlquist & Zona, 1988b), Pentaphylacaceae (1984c), Polemoniaceae (Carlquist et al., 1984), Scytopetalaceae (1988b), Stackhousiaceae (1987b), Staphyleaceae (Carlquist & Hoekman, 1985), Stilbaceae, including R etzia (1986), Sympetalae (1992b), Ticodendraceae (199 l a), Tovarioideae of Capparaceae (1985c), and Winteraceae (1989a). Similar comparative anatomical studies by others include the investigations of Alseuosmiaceae (Dickison, 1986c, 1989c), Araceae (French, 1987), Idiospermum of Calycanthaceae (Foreman, 1987), Caprifoliaceae and Adoxaceae (Ogata, 1988), Sarcandra of Chloranthaceae (Takahashi & Tamura, 1990), Coriaria (Suzuki & Yoda, 1989), Cyclanthaceae (Tomlinson & Wilder, 1984), mucilage cells in dicotyledons (Gregory & Baas, 1989), Didymelaceae (Takhtajan et al., 1986), Diegodendron (Dickison, 1988), Neowawraea in Euphorbiaceae (Hayden, 1987; Hayden & Brandt, 1984), Euphorbiaceae (Mennega, 1987), Neopringlea ofFlacourtiaceae (Lemke, 1987), Frankeniaceae (Whalen, 1987), Geosiris ofIridaceae (Goldblatt et al., 1984, 1987), Montiniaceae (Rakouth, 1989), Oceanopapaver (Schmid et al., 1984), Oleaceae (Baas et al., 1988), Penthoraceae (Haskins & Hayden, 1987), Darmera in Saxifragaceae (Gornail, 1989), Styracaceae (Dickison & Phend, 1985), Tetracarpaeaceae (Hils et al., 1988), Theaceae (Liang & Baas, 1990), and dicotyledons in general (Dickison, 1989b).

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7. Leaf Architecture
Comparative studies of the anatomy and architecture of leaves can also be highly enlightening as to the circumscription and relationships of the taxa examined. Among those recent investigations that I found especially informative are the publications on angiosperm laterocytic stomatotypes (Baranova, 1983), Alseuosmiaceae (Dickison, 1989c), Annonaceae (Stten & Koek-Noorman, 1986), Balanitaceae (Sarma & Rao, 1991), Barbeyaceae (Tobe & Takahashi, 1990), Bruniales (Carlquist, 1990c, 199 lb), Chloranthaceae (Todzia & Keating, 1988), Cunoniaceae (Dickison & Rutishauser, 1990), Cyclanthaceae (Wilder, 1984, 1985), Euphorbiaceae (Levin, 1986), Fagaceae (Jones, 1986), Hamamelidaceae (Li & Hickey, 1988; Pan et al., 1990), Hamamelididae (Wolfe, 1989), Lamiaceae (Cantino, 1990), Lilianae (Conover, 1991), Linaceae and relatives (Welzen & Baas, 1984), Melanthiaceae (Badawi, 1986), Menispermaceae (Wilkinson, 1989), Myrtales (Keating, 1984), Oleaceae (Inamdar et al., 1986), Philesiaceae (Arroyo & Leuenberger, 1988), Poaceae (Ellis, 1987; Judziewicz & Soderstrom, 1989; Renvoize, 1986a, 1986b, 1987a, 1987b), Polygalaceae (Baas, 1991; Detienne, 1991), Ranunculaceae (Hoot, 1991), Staphyleaceae (Dickison, 1987b), Tecophilaeaceae (Arroyo, 1986), Tepuianthaceae (Roth & Lindorf, 1990), Ticodendron (Hickey & Taylor, 1991), and Zosteraceae (Kuo et al., 1988).

8. Karyomorphology
Karyological investigations have been important to phylogenists since the early days of simple chromosome counting. Now, however, karyomorphology or karyosystematics has become a specialized branch of botany that involves not only the number and size of chromosomes but their morphology and internal anatomy as well. As thus circumscribed, karyosystematics is a relatively new scientific approach, and it offers much potential information for the clarification of relationships. Some of the recent groups that I think it has helped to elucidate are the following: Annonales s.s. and s.l., Malvales, and Violales (Morawetz, 1986a, 1986b, 1988), Araceae (Petersen, 1989), Aristolochiaceae (Morawetz, 1985), Asteranthos of Lecythidaceae (Kowal, 1989), Brunelliaceae and Caryocaraceae (Ehrendorfer et al., 1984), Apostasioideae of Orchidaceae (Okada, 1988), Piperaceae and Saururaceae (Okada, 1986), Sarcolaenaceae (Goldblatt, 1986a), Saxifragaceae s.s. (Soltis, 1988), Strelitziaceae (Manning, 1989), Styracaceae (Morawetz, 1991), Triuridaceae (Davidse & Martinez, 1990), and Walleria of the Tecophilaeaceae (Goldblatt & Manning, 1989). A number of publications presenting noteworthy chromosome records should be mentioned, as for New Caledonian plants (Carr, 1986) and Chinese endemics (Li & Hsu, 1986).

9. Micromorphology
Another relatively new scientific approach to the study of angiosperm relationships is that of micromorphology. It too offers new and often critical information for the phylogenists. Behnke has been one of the pioneers of this approach with his exhaustive study of sieve-element plastids throughout the angiosperms. Some of his more recent contributions have been studies of Acanthaceae (1986a), Annonales s.1. (1988a), Austrobaileyaceae (1986b), Buxaceae and Simmondsiaceae (1982), Dicotyledoneae (1991 a), Gunneraceae (1986c), Hamamelididae (I 989), Macarthuria of Mollugina-

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ceae (Behnke et al., 1983), Myristicaceae (1991 c), Myrtales (1984), Rhizophoraceae and Anisophylleaceae (1988b), Ticodendraceae (1991 b), and Winteraceae (Behnke & Kiritsis, 1983). The study of epicuticular waxes by Barthlott and his associates also offers new information of great interest in the Caryophyllales (Engel & Barthlott, 1988), Rosales s.l. (Fehrenbach & Barthlott, 1988), monocotyledons (Barthlott & Froelich, 1983; Froelich & Barthlott, 1988), and especially the Commelinanae (Froelich & Barthlott, 1987). Other studies involved myrosin cells and other protein-rich cells, as those with dilated cisternae (Jorgensen, 1987), ultrastructure of microhairs in Poaceae (Amarasinghe & Watson, 1988), morphology of orbicules (Ubisch bodies) in anthers of Chloanthaceae (Raj & E1-Ghazaly, 1987), and ultrastructure of nuclear inclusions in Verbenaceae and Oleaceae (Bigazzi, 1989).

10. Chemotaxonomy and Molecular Taxonomy


Perhaps the most promising comparative investigations for solving our problems of circumscription and searching out of natural relationships based on common origin are the various chemotaxonomic and molecular taxonomic approaches, especially the relatively new and currently very popular methods for analyzing the distribution of major structural rearrangements, as inversions and loss or gain of genes, as well as sequence variation, in chloroplast DNA (Palmer et al., 1988). Although in its infancy, this approach shows great promise. Some of the most informative recent studies are those by Beckstrom-Sternberg (1988) and Giannasi et al. (1992) on Caryophyllales, Bremer and Jansen (1991) on Rubiaceae, Downie and Palmer (1990) and Olmstead et ah (1992) on Asteridae, Givnish et al. (1990) on bromeliad subfamilies, Jansen and associates on Asteraceae (Jansen, 1990; Jansen & Palmer, 1988; Jansen et al., 1988, 1991; Keeley & Jansen, 1991; Kim el al., 1990; Michaels & Palmer, 1990), Olmstead and Palmer (1990) on Solanaceae, Ranker et al. (1990) on Bromeliaceae, Rettig et al. (1990) on Caryophyllidae, Soltis et al. (1990) on Saxifragaceae, Waterman (1990) on Rutaceae, Zhou and Jiang (1990) on Hamamelididae, and Martin and Dowd (1990) on the dicotyledons in general and legumes and proteads in particular. Timely warnings that molecular systematics "may represent an insidious new form of one-character taxonomy" and that gene trees may be quite incongruent with species trees have been offered by J. J. Doyle (1992) and Rieseberg and Soltis (1991). In general, the comparative biochemists have continued to offer much information of value to the phylogenists. Among the more recent papers offering significant information on the angiosperms generally are the publications by Ettlinger (1987) on glucosinolates, Gershenson and Mabry (1983) on secondary metabolites, Giannasi (1988) on flavonoids, Kubitzki and Gottlieb (1984) on micromolecular patterns, Molgaard (1985) on caffeic acid, Molgaard and Ravn (1988) on caffeoyl esters, Okuda (1987) on tannins, and Williams and Harborne (1988) on flavonoids in the monocots. More restricted as to target taxa are the recent studies on iridoids and quaternary amines in Acanthaceae (Jensen et al., 1988); phytochemistry recommending removal of Gisekia from Phytolaccaceae to Aizoaceae (Narayana & Narayana, 1988); mannose-binding lectins in Alliaceae and Amaryllidaceae (Van Damme et at., 1991); flavonoids in Alzateaceae (Graham & Averett, 1984); biflavonoids in "Julianiales" of the Rhoeae of Anacardiaceae (Wannan & Quinn, 1988) and in the wider Ana-

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cardiaceae (Young & Aist, 1987); neolignans and benzylisoquinolines in the Magnolianae (Gottlieb et al., 1989); phytochemical similarities of Apiaceae and Asteraceae (Holub et al., 1987); mannan in endosperm cell walls of Asteraceae but not in Dipsacales (Grau & Hopf, 1985); flavonoids in Balanitaeeae (Maksoud & E1 Hadidi, 1988); glucosinolates in Bretschneideraceae (Boufford et al., 1989); chemistry of Capparaceae (Anuradha et al., 1988); flavonoids in Cephalotaceae (Nicholls et al., 1985); sterols in Caryophyllales and putatively related taxa (Patterson & Xu, 1990); chiral esters in Eupomatia (Bergstrom et al., 1991); tigliane and related diterpenoids in Euphorbiaceae and Thymelaeaceae (Schmidt, 1987); phenolics in Gesneriaceae (Kvist & Pedersen, 1986); phytochemical profile and leaf phenolics in Gunneraceae (Doyle & Scogin, 1988a, 1988b); chemistry of "Hamamelididae" (Giannasi, 1986; Zhou & Jiang, 1990); flavonoids in Hydrangeaceae, Itea, and Saxifragaceae s.1. (Bohm et al., 1985, 1986, 1988); terpenoids in Icacinaceae (Kaplan et al., 1991); leaf flavonoids of Lactoris (Crawford et al., 1986); glycosides in Lamiaceae, Scrophulariaeeae, and related families (Tomas-Barberan et al., 1988); leaf flavonoids in Dahlgren's Liliiflorae (Williams et al., 1988); cyanogenic glycosides in Malesherbia and Passifloraceae (Spencer & Seigler, 1985; Spencer, 1987); liminoids in Meliaceae (Das et al., 1984); flavonoids in ericaceous Monotropoideae (Bohm & Averett, 1989); amino acid sequences in the Myrtales and other dicots (Martin & Dowd, 1986, 1988, 199 l); phenolic and amino acids and secondary metabolites of Averrhoa and the rest of Oxalidaceae (Devi & Narayana, 1990); paeoniitorin and other chemistry of Paeoniaceae (Yu & Xiao, 1987); fatty acids and seed oils in Pittosporaeeae (Stuhlfauth et al., 1985); sterols of Plumbaginaceae and Polygonaceae different from those in Caryophyllales (Wolfe et al., 1989); protein amino acid profiles of grass leaves and caryopses (Yeoh & Watson, 1987); chromones and lack ofliminoids in Ptaeroxylaceae (White, 1986); fiavonoids in Rhynchocalycaceae(Averett & Graham, 1984); iridoid glycosides in Rubiaceae (Inouye et al., 1988); chemosystematics of Rutaceae and of the rutaceous Dictyolomatoideae (Silva et al., 1988; Vieira et al., 1988), and phytochemistry of Sapindaceae and related families (Umadevi & Daniel, 199 l; Umadevi et al., 1986). 11. Phytoserology Phytoserological investigations continue to offer clues as to relationships of the taxa investigated. I remain somewhat skeptical of these comparative serological studies, even when they agree with my phylogeny, because they seem generally to agree with the phylogenetic trees currently most popular (but later often found less than realistic). However, careful serological studies do give us much to think about, as the following: legumin analysis in "Magnoliidae" and in dicots generally (Jensen, 1991; Jensen & Greven, 1984); analysis of nabisco and legumin in giucosinolateproducing dicots (Gerstberger, 1987); appraisal of"Amentiferae" (Petersen & Fairbrothers, 1985); investigations in Magnolianae (Fairbrothers & Petersen, 1983), Euphorbiaceae (Vogel, 1986), Juglandaceae (Polechko & Clarkson, 1986), and Liliales s.1. (Chupov, 1987; Chupov & Kutjavina, 1981); immunological relationships among subfamilies of Poaceae (Esen & Hilu, 1989, 1991; Hilu & Esen, 1988); serotaxonomy of Solanaceae (Lester & Roberts, 1986); and phytoserology of Typhales (Bergner & Jensen, 1989). Dahlgren (1983) stressed the importance of modem serological research in angiosperm classification.

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12. Paleobotany
a. Origins and Early Evolution of Angiosperms Although the paleontological studies in the Angiospermae usually do not clarify relationships among taxa, they frequently do indicate the time of origin of various major taxa, the early appearance of characteristics, habitat occupation, and pollination and dispersal syndromes, and thereby the direction of evolutionary trends. There lately have been numerous publications pushing the origins of many flowering plant taxa well back into Cretaceous or early Tertiary time. Evidence of members of the Magnoliales from mid-Cretaceous time has been presented as monosulcate (Early Aptian of Delaware and Virginia) and other pollen grains resembling those of Magnoliaceae, Chloranthaceae, Winteraceae, and Myristicaceae (Upper Barremian to Lowermost Aptian and Middle Albian of Israel) (Brenner, 1987; Ward et al., 1988); multifollicular fruit from the Mid-Cretaceous Dakota of Kansas (Crane & Dilcher, 1984; Dilcher & Crane, 1984); a floral axis bearing conduplicate carpels (from the Mid-Cretaceous of Japan) like those in extant Monimiaceae and Austrobaileya(Nishida, 1985); a possible magnolioid floral axis and magnoliaceous wood from the Upper Cretaceous of California (Page, 1984); lactorid pollen from boreholes off the southwestern coast of southern Africa (Zavada & Benson, 1987); and lauraceous flowers from the Mid-Cretaceous Potomac Group of eastern North America (Drinnan et al., 1990). Paleobotanical evidence for the early radiation of Chloranthaceae is especially strong with dispersed pollen, pollen in fossil androecia, and fossil leaves with chloranthoid teeth, architecture, and stomata (Crane, 1989a; Friis et al., 1986; Upchurch, 1988). Other Cretaceous taxa included leaves referred to the Trochodendrales from the Albian and Cenomanian (Crane, 1989a); platanaceous triaperturate pollen, inflorescences, flowers, and foliage from the Late Albian Patapsco Formation and Early Cenomanian (Crane, 1989b; Crane et al., 1986; Friis & Crane, 1989; Friis et al., 1988); buxaceous flowers and pollen from the North American Mid-Cretaceous (Drinnan et al., 1991); nothofagaceous pollen and leaves by the Santonian (Romero, 1986b); wood resembling that of extant Apocynaceae, Euphorbiaceae or Flacourtiaceae, Fagaceae, and Icacinaceae (Wheeler, 1987); caesalpinoid pollen (Crepet & Taylor, 1985); possibly portulacaceous pollen from the Iowa Cenomanian (Ravn, 1987); pollen of Rhoipteleaceae from the Maestrictian (Crepet, 1981); rutaceous Rutaspermum from the Late Cretaceous of East Germany, and an escallonioid flower from the Late Cretaceous of Sweden (Friis, 1990); gunneraceous pollen (as Tricolpites) first from Turonian with spread to high latitudes by Campanian (Jarzen & Dettmann, 1989); and Ulmaceae from the Late Cretaceous (Manchester, 1989a). Several excellent papers on the origins and early evolution of the angiosperms have been published by Basinger and Dilcher (1984), Brenner (1990), Muller (1984), Taylor and Hickey (1990), Tiffney (1984), Walker and Walker (1984), Friis (1984, 1989), Manchester (1987a), and Crepet et al. (1991). The Nebraska Dakota Formation, some 94 million years old (Ma), produced bisexual, actinomorphic, hypogynous, pentamerous, medium-sized flowers with clearly differentiated floral parts, petals alternate with sepals, stamens borne opposite the petals and producing small, tricolporate pollen grains, carpels fused, and receptacle disk present (Basinger & Dilcher, 1984). The flowers seem well adapted to insect pollination. Based upon Upper Cretaceous fossilized reproductive organs from Swedish formations, Friis (1984) hy-

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pothesized that epigyny and unisexual flowers developed in middle Cretaceous time or as early as the Cenomanian. Friis (1989) noted the appearance of syncarpous ovaries by mid-Cretaceous and of sympetalous corollas and probably zygomorphic flowers in Late Cretaceous time. Friis (1989) considered the earliest angiosperms as probably insect pollinated and generalists possibly pollinated by beetles, flies, micropterigid moths, sawflies and sphecid wasps. Adaptations to the more specialist insect groups were probably evolved during Mid- and Late Cretaceous. According to Crepet (1984) hymenopteran and lepidopteran pollinators and angiospermous taxa having flowers adapted to them were in existence at the time of major flowering plant radiation during the Upper Cretaceous. The earliest documented fruits and seeds of Cenomanian age (115-95 Ma) were small, with most of the fruits dehiscent follicles or capsules. The first fleshy fruit known is from the Cenomanian (98 Ma). Fruits and seeds increased in size in later periods with very large diaspores appearing in Early Eocene time (52 Ma) (Tiffney, 1984). The earliest Cretaceous angiosperm leaves from southern South America are from the Aptian (113-119 Ma) as large, lobate, craspedromous, and dentate with ramified tertiary veins and random 4th order venation (Romero & Archangelsky, 1986). In their analysis of North American Cretaceous leaf and wood assemblages, Wolfe and Upehurch (1987) and Upehurch and Wolfe 0987) found leaves to be generally small, thick-textured, mostly pinnately-veined, entire-margined, and unlobed, with rounded to emarginate apices common, and drip tips and liana types uncommon at low and middle palaeolatitudes, indicating warm, subhumid, aseasonal climates. Woods lacked growth rings, also indicating little seasonality. By the Upper Cretaceous Campanian in northern Montana leaf fossils are diverse and appear to represent five of the Cronquist-Takhtajan subclasses (Crabtree, 1987). Although the first verifiable and angiospermous fossils have been found in strata no earlier than Early Cretaceous time, ultimately fossils of earlier Mesozoic age will probably be found. Cornet (1989) has found additional Late Triassic fossils of Sanmiguelia, from the Sunday Canyon locality in Texas, that he interprets as angiospermous and apocarpous, combining both monocot and dicot characteristics, and homologous with male and female flower-like units of an associated gnetalean-like plant. Crane earlier (1985) concluded that the angiosperms were a sister group to the Gnetales and also to the Bennettitales and Pentoxylon. In an "experimental" cladistic study Doyle and Donoghue (1986) came to the same conclusions, and considered that these groups formed an anthophyte clade related to Caytonia and the glossopterids but possibly also to other Mesozoic seed-fern taxa. The true ancestry of the angiosperms, however, still is uncertain though their heritage does appear most likely among the Mesozoic seed ferns. Upchurch and Wolfe (1987) regarded the earliest angiosperms to have been unimportant early successional elements in the regional vegetation. Their gradual rise to dominance over the ferns and gymnosperms during Mid- and Late Cretaceous time has been variously explained. Bakker (1986) believes that the low-feeding, herbivorous, Cretaceous dinosaurs, with their square snouts and pincerlike beaks, helped the angiosperms beat-out the slower growing, less adaptive gymnospermous competition. Bond (1989) also credits the faster angiospermous reproductive and vegetative growth rates, the better transport system in stem and foliage, and the advantages of insect pollination and seed dispersal by animals as the explanations for the rise to dominance by angiosperms under conditions of favorable climate and

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nutrient-rich soils. Midgley (1991), however, has suggested that plant-animal interactions have been overemphasized and competitive interactions too much overlooked in the Cretaceous rise of the angiosperms. Only in the higher latitudes and elevations or on nutrient-poor soils, where the growth of the angiospermous competitors was reduced, could the gymnosperms maintain their dominance. Retallack and Dilcher (198 l) hypothesized that some of the earliest angiosperms were probably woody, microphyllous plants of the rift valley system joining Africa and South America during the Early Cretaceous. Adapted to pioneering disturbed coastal environments, they became widespread during the great transgressions and regressions of epeiric seas during the Mid-Cretaceous. Crane and Lidgard (1989) documented through fossil pollen the diversification and migrations of angiosperms from low paleolatitudes in Mid-Cretaceous to higher paleolatitudes in later Cretaceous time. b. Tertiary Paleobotany The terminal Cretaceous events, although they apparently had little effect on Southern Hemisphere climate and vegetation (Upchurch & Askin, 1990), did in the Northern Hemisphere have a major impact on climates and vegetation (Upchurch & Wolfe, 1987; Wolfe, 1987). There is evidence for a mass-kill of vegetation with a subsequent period of vegetative recovery. At low and middle northern palaeolatitudes precipitation levels increased greatly, leading to the widespread development of multistratal rainforests in areas ofmegathermal climates. This development of widespread closedcanopy vegetation provided many new or expanded niches selecting for tall, buttressed trees, lianas, epiphytes, understory herbs, and large seeds with substantial nutrient reserves. In high-middle palaeolatitude, mesothermal areas the Paleocene vegetation was largely deciduous, due apparently to the heavy extinction of broadleaved evergreen species. There was widespread selection for plants of deciduous habit and great diversification of extant families of deciduous, wind-pollinated angiosperms such as the Juglandaceae and Betulaceae, probably due to seasonal dryness. Early Tertiary fossils are much more numerous than Cretaceous fossils and easier to associate with extant taxa. Among those so identified are numerous and diverse fossils of many amentiferous families by Middle Eocene (Crepet, 1981); fruits of anacardiaceous Spondieae from the Middle Eocene Claiborne Formation (Grote & Dilcher, 1987); fruits and seeds of lasioid Araceae and cabombaceous Nymphaeales from the Middle Eocene Princeton chert of British Columbia, with palm and lythraceous remains (Cevallos-Ferriz, 1987; Cevallos-Ferriz & Stockey, 1988); Saballike palm leaves from the Middle Eocene Allenby Formation of British Columbia (Erwin, 1987); an extinct genus ofbetuloid fruit from the Oligocene of western North America (Manchester & Crane, 1987); diversity in Betulaceae by Middle Eocene (Crane, 1989b); fossil fruits of Ceratophyllum from Paleocene and later Tertiary (Herendeen et al., 1990); Cercidiphyllum-like infructescences, fruits and seeds from the British early Tertiary (Crane, 1984; Crane & Stockey, 1986); Eocene pollen of Euphorbiaceae, Loranthaceae, Sapindaceae, and Sapotaceae from the Middle Eocene of Tennessee (Taylor, 1989); mimosoid flowers and bipinnate leaves from the early Tertiary of Tennessee and Kentucky (Crepet & Taylor, 1985, 1986; Herendeen & Dilcher, 1987, 1990a, 1990b); all four subfamilies of Fabaceae represented by Middle Eocene (Herendeen, 1990; Herendeen & Dilcher, 1990b); castaneoid and trigonobalanoid pistillate inflorescences and dispersed mature fruits from the Paleocene/Eocene

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boundary of western Tennessee (Crepet & Nixon, 1989a); other Tertiary fossils of Fagaceae (Crepet, 1989; Crepet & Nixon, 1989b); gentianaceous Lisianthus pollen from the Eocene of Panama (Graham, 1984); Liquidambaroideae of Hamamelidaceae from Paleocene through Neogene (Ferguson, 1989); diverse juglandaceous pollen and fruits from the Paleocene and later (Manchester, 1989b, 1991); lauraceous and malpighiaceous flowers from the Middle Eocene Claiborne Formation (Taylor, 1988; Taylor & Crepet, 1987); magnoliaceous Liriodendron fruiting structures, carpels and leaves from the Miocene Clarkia flora of Idaho (Baghai, 1988); Nothofagus macrofossils from the Tertiary of Tasmania and western Antarctica (Hill, 1991); anthoecia of Poaceae as earliest known grass fossils (Thomasson, 1987); rosaceous Chamaebatiaria-like foliage from the Paleogene of western North America (Wolfe & Wehr, 1988); abundant rutaceous fossils in European and American Tertiary (Gregor, 1989); a fossil Smilax from Eocene sediments in western Tennessee (Sun & Dilcher, 1988); twigs with attached foliage, fruits, and flowers of the ulmoid Cedrelospermum from the Early Tertiary of Utah and Colorado (Manchester, 1989a, 1989c); fossil endocarps and achenes of Urticales from early Tertiary onwards (CoUinson, 1989); vesselless Nordenskioldia of the Trochodendraceae in the early Eocene (Crane et al., 1991), also with Trochodendron and Cercidiphyllum in the Miocene of northwestern North America (Manchester et al., 1991); and the earliest remains of grasses from the Paleocene/Eocene Wilcox Formation in western Tennessee (Crepet & Feldman, 1991). Pollen comparable with Anacardiaceae, Aquifoliaceae, Gunneraceae, Loranthaceae, Myricaceae, Myrtaceae or Sapindaceae, and Proteaceae were recovered from a deep well in Eocene sediments (Romero & Castro, 1986); and pollen grains referred to Arecaceae, Casuarinaceae, Chloranthaceae, Combretaceae/Melastomataceae, Euphorbiaceae, Myricaceae, Myrtaceae, Podocarpaceae, Restionaceae, Rutaceae, Sapindaceae, Sarcolaenaceae, and Winteraceae were identified from the Miocene of the southwestern Cape Region of South Africa (Coetzee & Muller, 1984; Coetzee & Praglowski, 1988).

13, Phytogeography
A much neglected approach to angiosperm phylogeny is phytogeography. Useful clues to familial relationships can sometimes be obtained from the past and present distribution of taxa (Thorne, 1989a). Perhaps the most significant recent phytogeographic publication (Krutzsch, 1989) analyzes paleochorologically a dozen phyletically important angiosperm groups. His analysis should serve as a vivid warning that present distributions may not at all be indicative of past distributions, especially in such taxa as Alangium, Ascarina, Ctenolophon, Nepenthes, Nypa, and Restionaceae. Many other anglospermous taxa known from the fossil record in North America no longer occur on that continent, at least north of Mexico; among them are Bombacaceae, Caldesia, Cercidiphyllum, Cleyera, DiUeniaceae, Engelhardtia, Eucommia, Euodia, Eupomatia, Exbucklandia, Gunnera, Icacinaceae, Mesoneuron, Phellodendron, Platycarya, Proteaceae, Pterocarya, Sabiaceae, Trapa, Winteraceae, and Zelkova (Thorne, 1986c; Tiffney, 1985; Wolfe, 1981). Numerous other archaic or at least relict taxa are represented today in North America by one or two species, such as Annona, Buckleya, Calycanthus, Clethra, Gordonia, Illicium, Liquidambar, Liriodendron, Menispermum, Nelumbo, Paeonia, Schisandra, and Syrnplocos. An even longer list surely could be prepared of extant taxa no longer present in Europe. Africa

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too has apparently been subject to extinction of numerous taxa, also presumably in Neogene time due to catastrophic climatic changes. Some of the recent phytogeographic publications of considerable interest clarify the distribution of the Annonaceae (Kessler, 1987), Araceae (Li, 1987; Mayo & Croat, 1987), Cercideae of the caesalpinoid Fabaceae (Wunderlin et al., 1987), Chloranthaceae (Todzia, 1988), Chrysobalanaceae and Lecythidaceae (Prance & Moil, 1983), Dipterocarpaceae (Ashton, 1982), Euphorbiaceae (Webster, 1987b), Lennoaceae (Yatskievych & Mason, 1986), Loranthaceae and Viscaceae (Barlow, 1983), mangroves (Tomlinson, 1986), Myrtaceae, Proteaceae, and Restionaceae (Johnson & Briggs, 1981), Pittosporales s.l. (Thorne, 1989a), Tristichioideae of Podostemaceae (Cusset & Cusset, 1988), Ranunculaceae (Zimam & Keener, 1989), and the African dendroflora (Ehrendorfer, 1988). Palaeoclimates and geography are clarified for North America by Wolfe (1981), Greller and Rachele (1984), and Frederiksen (1989), for Central America by Graham (1985, 1987), for South America by Romero (1986a), and for southeastern Asia by Bande and Prakash (1986). Phytogeographical exploration is still turning up numerous new species and genera and occasionally new families previously unknown to botanical science. Some of these new taxa fill in phyletic gaps or otherwise clarify phylogenetic or phytogeographic problems. Among the new taxa recently discovered in the field, or occasionally in the herbarium, are the following, listed alphabetically, that I have found especially significant phyletically or geographically: Acanthogilia (Polemoniaceae) from Baja California (Day & Moran, 1986); Anacampseros (Portulacaceae) in Argentina (Cullen, 1984); Archiatriplex (Chenopodiaceae) from China (Chu, 1987); Balgoya (Polygalaceae) in New Caledonia (Morat & Meijden, 1991); Beiselia (Burseraceae) from Mexico (Forman, 1987; Forman et al., 1989); Bentleya (Pittosporaceae) in southern Western Australia (Crisp & Taylor, 1990); a neotropical species of the otherwise Asian Caryodaphnopsis (Lauraceae) (Werff& Richter, 1985); Ceuthostoma (Casuarinaceae) from Malesia (Johnson, 1988); Chiangiodendron (Flacourtiaceae), representing a new tribe in Mexico for the New World flora (Wendt, 1988); two monotypic trigonobalanoid genera (Fagaceae), Colombobalanus and Formanodendron, the former from Colombia and the latter from southeastern Asia (Nixon & Crepet, 1989); Dahlgrenodendron (Lauraceae) from Natal and Pondoland (Merwe et al., 1988); a second species of Degeneria from Fiji (Miller, 1988, 1989); Dianthoveus (Cyclanthaceae) from South America (Hammel & Wilder, 1989); the Chinese Diplopanaxstachyanthus recognized as a living representative of Mastixicarpum, a fossil mastixioid (Cornaceae) (Eyde & Qiuyun, 1990); Faika and Parakibara (Monimiaceae) from Malesia (Philipson, 1985); Gamanthera (Lauraceae) from Costa Rica (Werff & Endress, 1991); Haptanthus of uncertain affinity, probably representing a new family, from Honduras (Goldberg & Nelson, 1989); Humulopsis (Cannabaceae) from eastern Asia (Grudzinskaya, 1988); two new lythraceous genera, Koehneria from Madagascar and Lourtella from Peru (Graham et al., 1986, 1987); Koyamaea (Cyperaceae) from Venezuela and Brazil (Thomas & Davidse, 1989); Lacandonia (Triuridaceae) of Mexico, treated by the authors as a new family (Martinez & Ramos, 1989); Mespilus canescens from Arkansas, a new generic record for North America (Phipps, 1990); Noahdendron (Hamamelidaceae) of northeastern Queensland (Endress et al., 1985); Nogalia of the boraginaceous Heliotropoideae from Somaliland and Arabia (Verdcourt, 1988); Nyssa talamanca (Cornaceae) in Costa Rica and Panama (Hammel & Zamora, 1990); Panamanthus (Loranthaceae) from Panama (Kuijt, 199 l); Tico-

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dendron of the fagalean Ticodendraceae from Central America (Gomez-Llaurito & Gomez P., 1989, 1991; Hammel & Burger, 1991); Waltheranthus (Gyrostemonaceae) from Western Australia (Keighery, 1985); and Xerospiraea (Rosaceae) from Mexico (Henrickson, 1985). Some of the more exciting range extensions include Pelliciera from Caribbean coasts (Roth & Grijalva, 1991), and Canella in the Yucatan Peninsula
(collected by A. Sanders).

14. Host-Parasite Relationships


Another often neglected approach to phylogenetic clarification is host plant-parasitic or -phytophagic relationships. Numerous clues to similar phytochemistry, thus often to basic relationships of the host plants, are offered by butterflies and other moderately specific phytophagous insects, by rusts, smuts, and other parasitic fungi (Thorne, 1979). Some informative publications of this nature that have come to my attention recently concern the host plants of papilionid (Miller, 1987) and nymphalid butterflies (Ackery, 1988) and of African and Australian butterflies generally (Ackery, 1991); rust susceptibility in the Rosaceae s.1. (E1-Gazzar, 1981); rust fungi and host plant coevolution (Hart, 1988); and relationships among the Poaceae, Cyperaceae, and Juncaceae as reflected by their rusts and smuts (Savile, 1987, 1990).

15. Cladistics
A most useful technical tool for clarifying intrafamilial relationships is the currently popular cladistic approach. In clarifying interfamilial and interordinal relationships this approach has frequently left much to be desired, largely because of the subjectivity of outgroup and character selection and determination of character polarity, failure to recognize linkage of characteristics in syndromes, overemphasis on parsimony, and scorning of paraphyletic groups and plesiomorphic characteristics. All too often the cladists appear to be more concerned with the elegance of their methodology than with the relationships and "family trees" of the taxa they are investigating. Furthermore, some of the cladistic zealots have angered botanists by their arrogance and insulting reference to plant phylogenists as "intuitive," "narrative," and "authoritarian," while wrapping themselves in a cloak of objectivity and legitimacy. For example, according to Humphries and Chappill (1988), cladistic methodology has "changed systematics from an enterprise based on intuition to a legitimate branch of biological science based on rigorous analysis of empirical evidence." Despite such shortcomings and irritations, this technical approach to phylogeny has lately been much refined and should in the future offer much help to the dedicated phylogenist. Some of the more informative recent cladistic studies include those of Atwood (1984) on the cypripedioid Orchidaceae, Barabe and Forget on the Calloideae of Araceae (1987), Brown and Varadaraj an (1985) on the Caryophyllales, Cantino (1982) and Cantino and Sanders on the Lamiaceae (1986), Campbell and Kellogg (1987) and Kellogg and Campbell (1988) on the Poaceae, Kalkman (1988 ) on the Rosaceae, Les on the Ceratophyllaceae (1988a), Loconte (1987) and Loconte and Estes (1989a) on the Berberidales (Ranunculales), Robson and Stevens (1987) on the Bonnetiaceae-Clusiaceae-Hypericaceae, Rodman (1991b) on glucosinolate-producing plants, Schwarzwalder and Dilcher (1991) on Platanaceae, Smith and van Wyk (1991) on Alooideae ofAsphodelaceae, TurnbuU and McNeill (1987) on Nepenthes, and Hufford and Dickison on Cunoniaceae (1992).

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Somewhat less successful, in my opinion, have been cladistic studies on major clades of the angiosperms (Dahlgren & Bremer, 1985; Donoghue & Doyle, 1989); tribal relationships in the Asteraceae (Anderberg, 1991; Bremer, 1987); Nymphaeales s.1. (Ito, 1987), Lactoris (Lammers et al., 1986), myrosinase plants (Rodman, 1987, 1988), Caryophyllales (Humphries & Richardson, 1980; Rodman et al., 1984) [see Hershkovitz (1989) for a critique of the latter study and the answer by Rodman (1990)], Hamamelididae (Hufford & Crane, 1989; Zavada & Dilcher, 1986), families of the Lamiiflorae (Lu, 1990), net-veined Liliiflorae (Conran, 1989), Magnoliidae (Loconte & Stevenson, 1991), Orchidaceae (Burns-Balogh & Funk, 1986a, 1986b) (see Dressier, 1987), Poales/Restionales (Manning & Linder, 1990), and "Rosidae" (Hufford, 1990). Cladism has been recently defended vigorously by Donoghue and Cantino (1988) and by Humphries and Chappill (1988).

III. Classification of the Angiospermae


A. CHANGES MADE IN THE SYSTEM SINCE 1983

I. Dicotyledoneae
As a result of all these above listed studies and numerous monographs and revisions listed below, I have had to make numerous changes in my system of classification published in 1983, and even in the most recent updated synopsis in Aliso (Thorne, 1992). A major change in my phyletic philosophy has been my decision to narrow my family and ordinal gap concepts to bring my acceptance of family and ordinal limits more in line with those of current taxonomists, especially in such difficult groups as my former broadly conceived Phytolaccaceae, Aizoaceae, Theaceae, Urticaceae, Linaceae, Rhizophoraceae, Saxifragaceae, Cornaceae, Araliaceae, Solanaceae, Loganiaceae, Verbenaceae, and Liliaceae. It should be noted, however, that my classification is still somewhat more conservative than that of most of my phylogenetic peers. Another important change has been my acceptance of the ending "-anae" for superorders instead of the traditional "-iflorae." This is in line with the treatments by Takhtajan, Cronquist, and the Dahlgrens, and is based on the criticism that logically "-iflorae" really is appropriate only for the flower-bearing Angiospermae. Also in the spirit of cooperation and compromise, I have decided to accept such widely used names as Magnoliopsida, Magnolianae, Magnoliales, Magnoliineae, Caryophyllanae, Caryophyllales, etc. Other names have been changed based on the principle of priority, despite the lack of a rule demanding priority for taxa above the level of the family. Dr. James Reveal (pets. comm.) has been most helpful in supplying dates for valid publication of ordinal and superordinal names. I have continued to use Lindley's Nixus Plantarum (1833) as my point of departure for ordinal names because of his consistent use of the "-ales" ending. Among other considerable changes in my system of classification are the deletion of Aristolochiineae but recognition of Austrobaileyineae, Sparattanthelioideae, and Ceratophyllales in Magnolianae (my former Annonanae). In Caryophyllanae, Phytolaccineae and Caryophyllineae are recognized. In Theanae significant changes include the deletion of Icacinineae with transfer of Icacinaceae and Cardiopteridaceae from Theales to Cornales; transfer ofDiegodendraceae (Rhopalocarpaceae, Spherosepalaceae) from Theales to Malvales; and transfer of Chrysobalanaceae from Rosales

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to Theales. Fouquieriineae have been elevated to Fouquieriales and transferred to Theanae. Celastrales are separated from Santalanae and elevated to Celastranae. Within Malvanae Cistineae are transferred back to Violanae from Malvales on a karyological basis, though they do seem rather intermediate between the two superorders. Sterculiineae and Malvineae and families Monotaceae and Diegodendraceae are recognized in Malvales; Gonystylaceae are recognized in and Aextoxicaceae removed from Euphorbiales (to taxa incertae sedis). In Violanae the suborder Salicineae is deleted with removal of Salicaceae to near proximity with Flacourtiaceae; Fusispermoideae are recognized in Violaceae; and Cucurbitineae are combined with Begoniineae. Moringaceae are removed from Brassicales and transferred to Sapindineae; Gyrostemonaceae are transferred from Sapindineae to Brassicales, likewise Salvadoraceae are transferred there from Oleales. Bataceae as Batales are also removed from Sapindineae and placed in Violanae after Brassicales. Geranianae are reordered with elevation of suborders Linineae, Geraniineae, and Polygalineae to ordinal rank and with addition of Rhizophorales. In Rutanae Juglandineae and Myricineae are removed and elevated to ordinal rank in Rosanae as Juglandales. Other prominent changes in the Rutales (including Sapindales) are the recognition in Sapindaceae of the subfamilies Koelreuterioideae, Hippocastanoideae, Aceroideae, and the removal of Gyrostemonaceae and Bataceae from the Sapindineae to the Violanae, as mentioned above. The combining of Hamamelidanae (Hamamelidiflorae) with Rosanae and the consequent transfer of Hamamelidales, Casuarinales, and Fagales and addition of Juglandales rather completely reorders Rosanae, along with recognition of Balanopales (my former Buxales), Bruniales, Saxifragales, Podostemales, and Cunoniales as separate orders in that superorder. Cunoniales are restructured with the reduction of the closely related Baueraceae, Eucryphiaceae, and Brunelliaceae to subfamily status in the Cunoniaceae. The rather closely related Cornanae are also rather thoroughly reorganized with recognition and inclusion of Hydrangeales; removal of Rhizophoraceae and Hippuridaceae from and elevation of Gunneraceae to Gunnerineae in Cornales; restructuring of Cornaceae--Nyssaceae; and addition to Cornales of Eucommiaceae (from Hamamelidales) and Icacinaceae and Cardiopteridaceae (from Theales). Pittosporales are transferred from the Rosanae to the Cornanae near the Araliales. In the Araliales the Araliaceae-Hydrocotylaceae-Apiaceae complex is treated as three separate families; and Aralidiaceae, Triplostegiaceae, and Morinaceae are recognized in the superorder. Calyceraceae are transferred from Dipsacales to Asterales of the Asteranae. Campanulales are transferred from Solananae to Asteranae, Sphenocleaceae are recognized, and Menyanthaceae are added from Gentianales. Brunonioideae are removed from the Goodeniaceae and transferred as Brunoniaceae to taxa incertae sedis. In Asteraceae the tribes Barnadesieae, Tarchonantheae, and Coreopsideae are accepted. In Solananae Fouquieriineae are removed as Fouqueriales to Theanae, and Hoplestigmataceae and Tetrachondraceae are added to the Boraginineae. Within Gentiananae the Oleales are deleted with Salvadoraceae removed to Brassicales and Oleaceae to Scrophulariineae. Loganiaceae are much trimmed down with removal of Desfontainia to Hydrangeales as Desfontainiaceae and Retzia to Stilbaceae in Scrophulariales adjacent to Buddlejaceae. Menyanthaceae are transferred from Gentiales (Rubiales) to Campanulales. In Scrophulariales (my former Bignoniales) Oleaceae and Stilbaceae join Buddlejaceae; Globulariaceae are recognized as a distinct

CLASSIFICATION AND GEOGRAPHY OF THE FLOWERING PLANTS

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family; Schlegelieae (Paulownieae) are transferred from Bignoniaceae to Scrophulariaceae; and CaUitrichaceae and Hippuridaceae are transferred from Lamiales and Cornanae respectively. Lamiales are reduced in rank to Lamiineae in Scrophulariales and Verbenaceae thoroughly disintegrated with removal of several groups, resulting in the recognition of Chloanthaceae, Avicenniaceae, Phrymataceae, Symphoremataceae, Nesogenaceae, and Tetrachondraceae as distinct families. Verbenaceae s.s. and Lamiaceae are completely reorganized with the help of Phillip Cantino (pers. comm.), with Verbenaceae essentially reduced to the former Verbenoideae, and the Lamiaceae restructured with the recognition of the Viticoideae, Ajugoideae, Scutellarioideae, Pogostemonoideae, Nepetoideae, and Lamioideae.

2. Monocotyledoneae
The subclass Monocotyledoneae has also been considerably reorganized, especially the Lilianae and Commelinanae. Within the superorder Lilianae I have recognized five orders: Liliales, Burmanniales, Asparagales, Dioscoreales, and Orchidales. The Liliales are divided into the suborders Melanthiineae, Liliineae, and Iridineae, and the Asparagales into the suborders Asparagineae and Amaryllidineae. I still regard the Melanthiaceae as the most archaic and least specialized monocots; whereas, I consider the Dioscoreales as relatively specialized within the Lilianae. Obviously, my formerly very complex and polyphyletic family Liliaceae has been largely dismantled, with recognition of most of the former subfamilies as distinct families in at least two orders, much as in the treatment by Dahlgren et al. (1985). The small Australasian family Hydatellaceae remains difficult to interpret but tentatively has been given its own order and superorder and placed between the Lilianae and Triuridanae. The Najadales and Najadaceae have been deleted from the Alismatanae with transfer of Najas to the Hydrocharitaceae. The related superorder Aranae has been changed considerably with the removal of the Acoraceae from the Arales to taxa incertae sedis and the complete reorganization of the Araceae based on the systems of Grayum (1990) and Bogner and Nicolson (1991). The superorder Typhanae has been deleted and the Typhales with single family Typhaceae, including Sparganioideae, placed tentatively in the Commelinanae between the Bromeliales and Zingiberales. My former broadly conceived order Commelinales has been redefined, with division into Bromeliales, Philydrales, Commelinales s.s., Juncales, and Poales. Velloziaceae, formerly treated near Bromeliaceae have been transferred to the Amaryllidineae, near the Hypoxidaceae, in the Asparagales. The Zingiberales are divided into the suborders Musineae, Strelitziineae, Lowiineae, Heliconiineae, Zingiberineae, and Marantineae; Commelinales into Xyridineae, Commelinineae, and Eriocaulineae; and Poales into HageUariineae and Poineae. In the FlageUariineae Joinvillea and Ecdeiocolea are elevated to family rank. It seems fitting that our most economically important and highly diverse, prolific, and specialized family Poaceae should sit atop the monocot "family tree." As usual, I have terminated the classification with a short list of Taxa Incertae Sedis to list therein those taxa whose position remains most uncertain. It is a shifting list that probably should be much longer than it is, and surely will be longer when all the anomalous genera are pruned away from the families where they are now misplaced.

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B. E X P L A N A T I O N OF THE PHYLETIC SHRUB

The cladists notwithstanding, I do not think it possible to erect an angiospermous "family tree." Although the Angiospermae are surely monophyletic, their two subclasses must have diverged from common seed-fern ancestors during the apparent Early Cretaceous or possibly late Jurassic origins of the class. Similarly, most of the superorders diverged early from their long extinct protoangiospermous ancestors. To draw one superorder or other major angiospermous taxon from another extant major taxon would be a futile exercise and most misleading. Hence, I prefer to treat the extant Angiospermae diagrammatically as a "shrub" diverging as superorders from a hollow center representing the extinct Protoangiospermae. The position of these superordinal stem cross-sections of the phyletic shrub indicates as closely as possible my interpretation of their interrelationships and their relative degree of specialization from their more primitive, archaic ancestors. Those farthest from the hollow center are the most specialized, those closest to the center the least specialized. The size of the superordinal balloons and the contained orders and suborders indicates approximately the number of species accepted for each major taxon, although the size of some of the smallest superorders has been exaggerated to make them more visible. Within the superordinal balloons the orders are shown as entire ellipses, the suborders as connected branches. This phyletic shrub replaces the one published in the Nordic Journal of Botany (Thorne, 1983) and differs from it rather considerably due to the major changes in my system of classification listed above. This diagram was drawn and lettered by my associates Oscar Dorado and Scott Zona at the Rancho Santa Ana Botanic Garden. Any errors are my own and any divergences from my classification are due to the continuing evolution of the latter. Unfortunately, any drawing will be somewhat obsolete by the time it is printed due to our rapidly expanding knowledge of the Angiospermae.
C. E X P L A N A T I O N OF THE SYNOPSIS

I. Philosophy of Classification
In the following synopsis of my classification of the Angiospermae I have continued to carry the hierarchy of the class down to the subfamily level where appropriate, and even down to the tribal level in the huge family Asteraceae. Subfamilies are important for they display intrafamilial divergence as well as their immediate common ancestry with other subfamilies in the same family. Despite my narrowed family gap concept, I still prefer not to multiply taxa unnecessarily where common recent ancestry seems evident. Thus I would rather not break up such well defined, if very large, families as the Papaveraceae, Fabaceae, and Apocynaceae whose subfamilies are closely linked by their characteristics and by intermediate taxa. Other taxonomists, however, do object to my extended hierarchy, and often treat my subfamilies as distinct families. The difference in ranking of taxa is not important so long as the relationships are firmly based on all pertinent data. Anyone interested in my philosophy of classification can find it outlined in my Evolutionary Biology paper (Thorne, 1976) and earlier publications (Thorne, 1958, 1963a, 1975). It has not changed greatly over the years, though I have, as mentioned above, seen fit to narrow somewhat my ordinal and family gap concepts, leading to

CLASSIFICATION AND G E O G R A P H Y OF THE FLOWERING PLANTS

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my acceptance of somewhat more narrowly defined orders and families. The changed total numbers of all these taxa down to the genus and species are listed in Table I. Because of space limitations I have not attempted here to give my reasons for the circumscriptions and alignments accepted in this classification. I am preparing a book which will outline all my reasons and cite the numerous studies, revisions, monographs, etc. from which I have taken my data and drawn my conclusions. In the meantime, one can find discussions of many of my realignments, especially those most divergent from my phylogenetic peers, in my previous phylogenetic publications (as Thorne, 1968, 1973a, 1974a, 1974b, 1974c, 1975, 1976, 1977a, 1978b, 1979, 1981, 1983, 1985, 1989a, 1989b; Thorne et al., 1977; Thorne & Scogin, 1978).

2. Nomenclature
Names used for the various taxa in the synopsis are according to the International Code of Botanical Nomenclature (Greuter et al., 1988), although I have extended the principle of priority to all categories up to the class, as mentioned above. Synonyms or additional included or excluded taxa are listed usually only where the names or treatment of taxa deviate considerably from those in A. Engler's Syllabus der Pflanzenfamilien, ed. 12, vol. 2 (Melchior, 1964). Subfamilial treatment is based largely upon those authorities that I regard as best informed and phyletically most realistic in their classification. A number of larger families still await realistic subdivision. In a few families well known to me I have devised my own tentative subfamilial classification.

3. Recent Monographs and Revisions


Some of the recent taxonomic treatments that I have found to be especially helpful in the preparation of this synopsis are those on the Acanthaceae (Barker, 1986), Aizoaceae (Bittrich & I-Iartmann, 1988; Bittrich & Struck, 1989), Alzateaceae (Graham, 1984; Tobe & Raven, 1984c), Araceae (Bogner & Nicolson, 1991; Grayum, 1990), Balanophoraceae (Hansen, 1986), Blepharocarya of Anacardiaceae (Wannan et al., 1987), Arecaceae (Dransfield et al., 1990; Uhl & Dransfield, 1987), Heliantheae ofAsteraceae (Eriksson, 1991), Berberidaceae (Loconte & Estes, 1989b), Brassicaceae and other glucosinolate-producing plants (Rodman, 1991a, 1991b), Bromeliaceae (Gilmartin & Brown, 1987; Gilmartin et al., 1989), Burmanniaceae (Maas et al., 1986), Buxaceae (Friis, 1989b), Cactaceae (Barthlott, 1988), Campanulales (Lammers, 1990), Casuarinaceae (Johnson, 1989), Ceratophyllaceae (Les, 1986, 1988c), Chrysobalanaceae (Prance & White, 1988), Commelinaceae (Faden & Hunt, 1988, 1991), Cucurbitaceae (Jeffrey, 1990), Cunoniaceae (Hufford & Dickison, 1992), Cynomorium (Lye, 1991), Cyperaceae (Goetghebeur, 1986), Daphniphyllaceae (Sutton, 1989b; Zhang & Lu, 1989), Didymelaceae (Sutton, 1989a), Ehretioideae of Boraginaceae (Miller, 1988), Eremolepidaceae (Kuijt, 1988), Euphorbiaceae (Webster, 1987a), Fagaceae (Nixon, 1989), Flacourtiaceae (Lemke, 1988), Gesneriaceae (Wiehler, 1983), Gnaphalieae of Asteraceae (Anderberg, 1991), Gyrostemonaceae (George, 1982), Haemodoraceae (Macfarlane et al., 1987; Simpson, 1990), Hamamelidaceae (Endress, 1989c), Heliantheae of Asteraceae (Eriksson, 1991), Heliconiaceae (Kress, 1990b), Iridaceae (Goldblatt, 1990), Juglandales (Stone, 1989; Lu & Zhang, 1990), Krameriaceae (Simpson, 1989), Lamiaceae/Verbenaceae (Cantino, 1990, 1992a, 1992b; Sanders & Cantino, 1984), Lardizabalaceae (Qin, 1989), Lauraceae (Werff,

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1991), Leitneriaceae (Jarvis, 1989), Lennoaceae (Yatskievych & Mason, 1986), Lilianae (Clifford et al., 1987; Conran, 1987; Conover, 1991; Matthew, 1988), Limnocharitoideae (Haynes & Holm-Nielsen, 1990), Maloideae of Rosaceae (Phipps et al., 1991; Robertson et al., 1991), Malpighiaceae (Anderson, 1990), Medusagynaceae (Robertson et al., 1989), Meliaceae (Cheek & Rakotozafy, 1991), Monimiaceae (Philipson, 1986a, 1987b, 1988), Monocotyledoneae in general (Dahlgren et al., 1985), Monotaceae (Kostermans, 1985), Morinaceae (Cannon & Cannon, 1984), Moraceae (Humphries & Blackmore, 1989), Myricaceae (MacDonald, 1989), Myrtaceae (Johnson & Briggs, 1984; Schmid, 1984; Tobe & Raven, 1990), Myrtales in general (Dahlgren & Thorne, 1984; Vliet & Baas, 1984), Nolanaceae (Mesa, 1981), Nothofagaceae (Hill & Read, 1991), Orchidaceae (Dressier, 1986, 1990a, 1990b), Pedaliaceae (Manning, 1991), Physenaceae (Takhtajan, 1985), Philydraceae (Adams, 1987), Poaceae and subfamilies (Barkworth & Everett, 1987; Clayton, 1987; Clayton & Renvoize, 1986; Conert, 1987; Crins, 1991; Jacobs, 1987; Macfarlane, 1987; Soderstrom & Ellis, 1988; Soderstrom et al., 1987), Polygalaceae (Meijden, 1982), Portulacaceae (Carolin, 1987; Hershkovitz, 1991), Ptaeroxylaceae (White, 1990), Ranunculaceae (Duncan & Keener, 1991; Fu, 1990; Hoot, 1991), Rhynchocalycaceae(Tobe & Raven, 1984b), Ripogonaceae (Conran & Clifford, 1985), Rubiaceae (Lorence, 1990), Scoliopeae of Liliaceae (Utech, 1992), Selaginaceae (Hartley & Balkwill, 1990), Sparganium of Typhaceae (Cook & Nicholls, 1986, 1987), Swartzioideae of Fabaceae (Ferguson & Skvarla, 1988), Thalictroideae (Fu, 1990), Trimeniaceae (Philipson, 1986b), Triplostegiaceae and Morinaceae (Hofmann & Gottmann, 1990), Tropaeolaceae (Sparre & Anderson, 1991), Urticaceae (Friis, 1989a), Urticales (Berg, 1989), Velloziaceae (Mello-Silva, 1991), Violaceae (Hekking, 1988), and Zingiberales (Kress, 1990a). 4. Symbols and Numbers Because the reader deserves some indication of the degree of confidence I place in the alignment used, hierarchical level assigned, circumscription accepted, or all of these, I have used in the synopsis for each category above the subfamily a simple "A," "B," "C" scale to indicate degree of confidence. "A," as used with Lactoridaceae, Chrysobalanaceae, and Ceratophyllales, represents limited confidence in the position of all three taxa and in addition the hierarchical ranking of the last taxon. Any less confidence would condemn a taxon to taxa incertae sedis. "B," as used with Paeoniales, Raittesianae, Paracryphiaceae, Oncothecaceae, Asteropeiaceae, Tetrameristaceae, etc., suggests that there is some evidence that the alignment, hierarchical ranking, or circumscription is probably correct. "C," used generally throughout the synopsis implies considerable confidence that accumulated data have allowed a realistic placement and circumscription. The numerals following most of the taxa listed are the number of genera and species (15/265 indicating 15 genera and 265 species) accepted for that taxon. Most were taken from Willis's Dictionary of the Flowering Plants and Ferns, Student Ed. (Shaw, 1985) or The Plant Book. A Portable Dictionary of the Higher Plants (Mabberley, 1987) except where more accurate information could be found in recent, reliable monographs and revisions. Some of these latter that should be cited are treatments of Alismataceae (Rogers, 1983), Apocynineae of Apocynaceae (Rosatti, 1989), Aponogetonaceae(Bruggen, 1985), Araliaceae (Philipson, 1979), Boraginaceae

CLASSIFICATION AND GEOGRAPHY OF THE FLOWERING PLANTS

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(AI-Shehbaz, 1991), Buxaceae (Koehler & Breuckner, 1989), Cactaceae (Hunt & Taylor for Consensus Committee, 1990), Campanulaceae and Sphenocleaceae (Rosatti, 1986), Commelinaceae (Tucker, 1989), Crossosomataceae (Holmgren, 1988), Dioscoreaceae (AI-Shebaz & Schubert, 1989), Elatinaceae (Tucker, 1986), Ericoideae (Oliver, 1989), Eriocaulaceae (Kral, 1989), Fumarioideae (Liden, 1986), Hydatellaceae (Cooke, 1987), Hydnoraceae (Musselman, 1989), Hypecoideae (Dahl, 1987), Leitneriaceae (Jarvis, 1989), Maloideae ofRosaceae (Phipps et al., 1990), Metteniusa (Lozano-C & Lozano, 1988), Moringaceae (Verdcourt, 1985), Nesogenaceae (Marais, 1981), Nothofagaceae (Philipson & Philipson, 1988), Orchidaceae (Atwood, 1986), Paniceae (Webster, 1988), Piperaceae (Bornstein, 1991), Pittosporaceae (Friis, 1987 ), Plantaginaceae (Rosatti, 1984), Pontederiaceae (Eckenwalder & Barrett, 1986), Portulacaceae (Nyananyo, 1990), Posidoniaceae (Kuo et al., 1990), Potamogetonaceae (Les & Sheridan, 1990; Wiegleb, 1988), Pteleocarpa (Veldkamp, 1988), Restionaceae (Linder, 1985), Sparganioideae (Cook & Nicholls, 1987), Stackhousiaceae (Barker, 1984), Tamaricaceae (Crins, 1989), Theophrastaceae (Stahl, 1991), Triuridaceae and other saprophytes (Maas & Rfibsamen, 1986), Xanthorrhoeaceae and Dasypogonaceae (Bedford et al., 1986), Xyridaceae (Kral, 1983), Zannichelliaceae (Haynes & HolmNielson, 1987), Zosteraceae (Kuo et al., 1988), plus the various monographs cited above. For the higher categories the larger numbers of species have been rounded offto the nearest five or ten to avoid spurious exactitude. Where a family or subfamily has less than four genera recognized, the genera are listed.

5. Distribution of Taxa
For each family, subfamily, and tribe in the synopsis the known distribution is listed, mostly in greatly abbreviated form, for the more widespread taxa. The abbreviations used are explained in Table V preceding the synopsis. The distributional information has been gleaned from recent monographs, floras, plant geography studies, and notes on range extensions too numerous to list here. Most helpful of all were the four published volumes of Pacific Plant Areas published by the Rijksherbarium at Leiden (1963-1984) and Plant-Geography of the Pacific by M. M. J. van Balgooy (1971). Other data have been gleaned from some of my own biogeographical publications (Thorne, 1963b, 1965, 1969a, 1969b, 1972, 1973b, 1977b, 1978a, 1986a, 1986b, 1986c). For some taxa herbarium specimens produced information not yet in print (as the presence of Canellaceae in Mexico). For both Old and New Worlds I have attempted to list the northernmost and southernmost limits of most ranges, preceded usually by an -, as N(--Green), indicating occurrence north to Greenland in North America, and Au(-Tas), indicating occurrence south in Australia to Tasmania. Lack of an -, as At(Ber, Mad), states that the taxon is reported only from Bermuda and Madeira of the Atlantic islands; whereas presence of the -, as Me(-Fiji) proclaims that the taxon is known to reach eastward in Malesia at least to the Fiji Islands, P(-Marq) that the taxon reaches eastward in the Pacific to the Marquesas Islands, or P(-Bon, Gal) that the taxon is known in the Pacific only eastward to the Bonin Islands and westward to the Galapagos Islands. The entire distributional pattern for each taxon should clarify the abbreviated listing. The geographical abbreviations n, ne, s, sw, e, w, and c, of course, should be interpreted as north, northeast, south, southwest, east, west, central, etc. All geographical abbreviations are decapitalized unless part of the country, state, or provincial name.

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6. Arrangement of Tara in Synopsis


It must be emphasized that each superorder in the synopsis is merely one line of evolution within the numerous-stemmed, many-branched complex phylogenetic shrub. A phylogenetic tree, given our limited paleontological record, is not realistic. A "phylogenetic" hedge, despite its double-entendre appeal, is also not realistic because I do firmly believe in the monophyletic origin of the Angiospermae and of their two subclasses, as well as most of the lesser-ranked taxa. We hope that most of the polyphyletic groupings have been rooted out of the classification but surely some remain to be extirpated, a probable example of such being the Lamiaceae as usually accepted (Cantino, 1990, 1992a). As each superorder, order, or suborder terminates, the classification drops back down the evolutionary ladder to the beginning of the next major line of ascent. By placement in the synopsis I have tried to indicate closeness of relationship and increasing specialization. A linear sequence cannot, of course, approximate the probable branchings nor indicate the numerous interrelationships among the superorders and lesser taxa. It is hoped that the phylogenetic shrub, illustrated by Figure 1, will be more helpful in these respects.

7. Discussion of Tables
In developing this synopsis of the Angiospermae a large amount of data was accumulated. In order to make it more readily available I have summarized the information in four tables. Table I presents a statistical summary of the various angiosperm taxa in the two subclasses: species, genera, subfamilies, families, suborders, orders, and superorders. The best estimate I could obtain for number of species in the angiosperms was 233,885, give or take a few thousand, with the dicots about three times more numerous than the monocots. In genera the monocots are less than one quarter of the 12,650 angiosperm genera, and in families they are about onefifth of the 437 families that I currently accept as valid. Because some of my phylogenetic peers (Cronquist, 1981; G. Dahlgren, 1989a, 1989b; R. Dahlgren et al., 1985; Takhtajan, 1987) accept many of my subfamilies as valid families, I have chosen to use subfamilies and undivided families as the most significant units. Thus the Angiospermae consist of 708 subfamilies and undivided families (or 437 families and 271 additional subfamilies, i.e., subfamilies other than the typical subfamilies). Since I have often been accused of being overly conservative in my recognition of families, I felt it might be informative to count the oligogeneric and oligotypic families listed in the synopsis, 217 of the former and 97 of the latter. Although it may be distressing to the student of flowering plants to have to deal with 48 monospecific families and 32 monospecific subfamilies for a total of 80, one should remember that the angiosperms have been evolving on earth for at least 120 million years and have suffered very heavy extinction even before modern man became too numerous and began to eliminate most of the species-rich environments. Such extinctions have broken up the near phyletic continuum and made it possible to develop our classifications, largely based upon phyletic gaps. Table II, also drawn from the Synopsis, summarizes the distribution about the world in the major regions of those families and subfamilies believed to be indigenous in each. Asia, even without Malesia, is by far the richest region phyletically, partly because of its latitudinal width from the Arctic nearly to the Equator, partly because

CLASSIFICATION A N D GEOGRAPHY OF THE FLOWERING PLANTS

249

cD 0

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250

THE BOTANICALREVIEW

Table I Statistical s u m m a r y o f the class Angiospermae (flowering plants) as interpreted from this synopsis
Dicotyledoneae Species Genera Subfamilies Families Subfamilies and undivided families Suborders Orders Suborders and undivided orders Superorders Monogeneric families Monogeneric subfamilies Monotypic families Monotypic subfamilies Digeneric families Digeneric subfamilies Ditypic families Ditypic subfamilies Trigeneric families Trigeneric subfamilies Tritypic families Tritypic subfamilies 175,690 9882 313 351 566 53 52 86 19 123 83 42 28 27 29 31 16 30 16 13 8 Monocotyledonae 58,195 2768 87 86 142 18 19 31 9 25 17 6 4 6 7 2 4 6 7 3 2 Total Angiospermae 233,885 12,650 400 437 708 71 71 117 28 148 100 48 32 33 36 33 20 36 23 16 10

Table II Putatively indigenous angiosperm families and additional subfamilies o f the world
Total families and add. subfamilies

Dicots Asia (excl. Malesia) Families Add. subfamilies 226 127

Monocots 61 27 57 32 46 32 54 27 53 24

Totals 287 154 270 141 240 132 246 123 246 107

441

South America (incl. Trinidad) Families 213 Add. subfamilies 109 Central America (incl. Mexico) Families 194 Add. subfamilies 100 Africa (S of Sahara) Families Add. subfamilies Malesia (Malaya-Fiji) Families Add. subfamilies 192 96 193 83

411

372

369

353

CLASSIFICATIONAND GEOGRAPHYOF THE FLOWERING PLANTS Table II Continued

251

Dicots Australia (incl. Tasmania) Families Add. subfamilies North America (N of Mexico) Families Add. subfamilies West Indies (incl. Bahamas) Families Add. subfamilies Madagascar & Comoros Families Add. subfamilies Pacific Basin (excl. Hawaii) Families Add. subfamilies New Caledonia & Loyalties Families Add. subfamilies Europe (incl. Med. Africa) Families Add. subfamilies Indian Ocean Islands Families Add. subfamilies Atlantic Islands Families Add. subfamilies New Zealand Families Add. subfamilies Hawaiian Islands Families Add. subfamilies Subantarctic Is. Families Add. subfamilies Antarctica Families Add. subfamilies 160 83 165 79 158 67 154 67 142 60 121 47 114 49 115 50 96 32 88 19 70 15 35 2 1 0

Monocots 60 29 42 23 36 25 40 23 32 22 33 15 35 13 31 16 27 8 24 7 13 5 10 3 1 0

Totals 220 112 207 102 194 92 194 90 174 82 154 62 149 62 146 66 123 40 112 26 83 20 45 5 2 0

Total families and add. subfamilies

332

309

286

284

256

216

211

212

163

138

103

50

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THE BOTANICALREVIEW

Table III Angiosperm families and subfamilies o f limited distribution


Total families and additional subfamilies 38 36 26 30 17 11 5 2 2 1 168

Major geographic units 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. Africa (mainland south of Sahara) South America (incl. Trinidad) Asia (excl. Malesia) Australia North America (incl. Central America and West Indies) Madagascar (and Comoro Is.) New Caledonia Malesia (to Fiji) Indian Ocean Pacific Basin Totals

Endemic families 21 20 17 14 10 7 5 2 1 1 98

Endemic subfamilies 18 16 10 16 7 5 0 0 1 0 73

it is the crossroads between Africa, Europe, and the Americas, and partly because o f its easy access to Australasia through immediately adjacent Malesia. It is also significant that the sometime insular continent o f South America is second richest in families and subfamilies because o f its latitudinal stretch from the Antarctic to north o f the Equator, its varied topography from the low-elevation equatorial rainforests along the A m a z o n to the snowfields o f the highest Andes, and its relatively recent junction with N o r t h America through the Panamanian Isthmus and earlier contacts with that continent via Antillean and Central American island stepping-stones. For-

Table IV W o r l d distribution o f angiosperm families and subfamilies


Subfamilies and undivided families 132 26 94 7 12 9 261 65 47 22 15 44 28 40 167 708

Families Subcosmopolitan Missing from one continent Pantropical Missing from one continent Widespread temperate Southern Hemisphere Other narrower disjuncts Eurasian (incl. Malesian)-American N. American (incl. C. Am. & W. Indies)-S. America American-African Australian-S. American Africa-Madagascar-Eurasia; -Australia; -Pacific Asia-Malesia; -Australia; -Pacific Other disjunctions Endemic to one continent Totals 97 18 49 5 9 7 154 35 21 15 7 24 13 39 98 437

Subfamilies 89 15 70 3 8 4 138 36 27 12 8 20 15 20 73 400

CLASSIFICATIONAND GEOGRAPHYOF THE FLOWERINGPLANTS

253

Table V Key to abbreviations for geographic areas in following synopsis


Antarc-Antarctic or subAntarctic in distribution; often ranging northward in the Andes and other high mountains Arct--Arctic in distribution; often also in alpine locations to the south Bor--Essentially boreal in distribution, i.e., distributed between the Arctic and temperate climatic areas Temp-Essentially or totally temperate in distribution TempMont--Primarily restricted to mountains in the temperate zone; basically Arctic-alpine plants Semitr--Essentially semitropical in distribution Trop--Essentially tropical in distribution TropMont--Primarily restricted to higher mountains in the tropics Subcosm-Widely distributed on all habitable continents and on many islands, i.e., nearly cosmopolitan (no angiosperm family being truly cosmopolitan in occupying all possible habitats) Af-Africa south of the Sahara Desert As-Asia: Arabian Peninsula and Anatolia east to Bering Strait incl. continental islands of Sri Lanka (Ceylon), Hainan, Taiwan, and Japan (Near East and all of Eurasia south of the Caucasus and east of the Ural Mountains) At--Atlantic Ocean islands, including Bermuda and Macaronesia south to Gough and Tristan da Cunha islands Au-Australia, including Tasmania and Lord Howe Island C--Central America, incl. Mexico, south through Panama E--Europe (north of the Black Sea and Caucasus Mountains and west of Ural Mountains) but incl. Mediterranean area and Saharan North Africa G--New Caledonia (Grande Terre) and Loyalty Islands H--Hawaiian Islands I--Indian Ocean islands, incl. Socotra, Aldabra, Seychelle, and Mascarene south to Amsterdam and St. Paul islands Ma--Madagascar and Comoro Islands Me--Malesia, a largely archipelagic area including the Malayan Peninsula on the west to the Fiji Islands on the east, including Malaysia, Indonesia, the Philippines, Papua New Guinea, Solomon Islands, Vanuatu (New Hebrides), and Fiji, or basically all the continental islands southeast of Asia, north of Australia, and southwest of the Pacific Basin N--North America N of Mexico, incl. Greenland and the Canadian Arctic Archipelago O--Subantarctic islands south of the Antarctic Convergence P--Pacific Ocean islands north and east of Malesia, New Caledonia, and New Zealand (all the oceanic islands of the Pacific Basin except the Hawaiian Islands; i.e., Bonins, Micronesia, Polynesia, Guadalupe, Revillagigedo, Cocos, Clipperton, Galapagos, and Juan Fernandez islands S--South America, including Netherland Antilles and Trinidad, south to and including Tierra Del Fuego and Falkland Islands Wind--West Indies (Bahama Islands, Greater and Lesser Antilles) Z--New Zealand and Chatham Islands Additional geographic abbreviations used in list Alas-Alaska, northwesternmost state of USA, mostly with boreal and Arctic flora except for temperate, largely insular southern panhandle extending south to Prince Rupert Sound north of the queen Charlotte Islands of British Columbia Alb-Alberta, Canadian province between British Columbia and Saskatchewan N of Montana, and S of District of Mackenzie Amaz-Basin of the Amazon River and tributaries, mostly in Brazil and Ecuador but including parts of Venezuela and Colombia south to Bolivia Ant--Antilles Arab--Arabian Peninsula, treated here as southwesternmost Asia, but southern tip with strong African floristic elements Arg--Argentina, with Chile, the southernmost republics of South America Az--Azores, a Portugese archipelago in eastern Atlantic

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Table V Continued
Balk--Balkans, several nations of southeastern Europe between Adriatic and Aegean and Black
seas

BCal--Baja California, Mexican peninsula south of California Bel--Belize, formerly British Honduras, on the Yucutan Peninsula, east of Guatemala and south of Mexico Ber--Bermuda, island nation in Atlantic Ocean east of North Carolina with a subtropical flora due to climatic influence of the adjacent Gulf Stream Bis--Bismarck Archipelago, islands between New Guinea and the Solomon Islands and south of the Caroline Islands Bol--Bolivia, South American nation west of Brazil and north of Paraguay, Argentina, and Chile, much of it in the Amazon Basin east of the Andes Bon--Bonin and Volcano Islands, in the western Pacific Ocean south of Japan and north of the Mariana Islands Born--Borneo, very large island between Sumatra and Philippines and west of Sulawesi (Celebes) Braz--Brazil, large South American nation south of Guayana and Venezuela and east of Peru and Bolivia BrC--British Columbia, southwesternmost province of Canada, N of Washington, E of S Alaska, S of Yukon Territory and Mackenzie District, and W of Alberta Cal--California, W USA Can-Canary Islands, part of Macaronesia, Spanish islands in Atlantic west of Morocco and south of Madeira Cape--Cape Peninsula at tip of southern Africa CapeP-Cape Province, southernmost province of Republic of South Africa CapeV--Cape Verde Islands, part of Macaronesia, formerly Portuguese islands in Atlantic west of Senegal on the Bulge of western Africa Car-Caroline Islands, including Palau and Yap, in southwestern Micronesia, north of New Guinea and Bismarck Archipelago and south of Mariana Islands Cauc--Caucasus Mts., range of mountains between Black and Caspian seas, separating southern Europe from SW Asia Cey--Ceylon Chi--China Col--Colombia, northwesternmost republic of South America CRica--Costa Rica, Central American republic north of Panama and south of Nicaragua Desv--Desventuradas, Chilean islands of San Felix and San Ambrosio, in Pacific west of northern Chile East--Easter Island, Chilean island in southern Pacific Ocean ese of Polynesia Ecu--Ecuador, western South American nation on equator, lying between Colombia and Peru Egy--Egypt, northeasternmost nation of Africa, incl. Sinai of SW Asia Eth--Ethiopia Fern-Juan Fernandez Islands, the Chilean islands Mas a Tierra and Mas a Fuera, in Pacific west of Santiago, Chile Ha--Florida, peninsular state south of Georgia, southern tip of which, with the adjacent Florida Keys, has an essentially Caribbean flora Fuegia--Tierra del Fuego, archipelago belonging to Chile and Argentina, south of Strait of Magellan Gal--Galapagos Islands, oceanic, volcanic islands in Pacific on the equator, west of and belonging to Ecuador Green--Greenland, large island northeast of North America, now with interior largely covered by deep glaciers Guad-Guadalupe Island, oceanic, high volcanic Mexican island in Pacific off coast of Baja California GuayH--Guayana Highlands, sandstone table mountains (tepuis) in Guyana, Venezuela, and northern Brazil, with rich and possibly archaic flora Guy-Guyana, formerly British Guiana, small nation between Venezuela and Surinam and north of Brazil

CLASSIFICATIONAND GEOGRAPHYOF THE FLOWERINGPLANTS

255

Table V
Continued Him--Himalayas, extensive high mountain ranges north of India Ice-Iceland, Scandinavian island republic in north Atlantic between Greenland and Norway and northwest of the British Isles Ire--Ireland, westernmost large continental island of the British Isles Jam-Jamaica, large Antillean island south of Cuba and west of Hispaniola Jap-Japan, temperate continental archipelago east of Russia and Korea Kam--Kamchatka Peninsula of the Russian Far East north of Kurile Islands and west of Bering Sea and Aleutian Islands; here used to include Chukchi Peninsula and other extreme northeastern parts of Russia Kor-Asiatic peninsula between Yellow and Japanese seas, south of Manchuria, Korea, and southwest of the Russian Far East La--Louisiana, SE USA Lab-Labrador, northeasternmost part of Canadian mainland, NW of Newfoundland and part of Newfoundland Province Mac--Macaronesia, several Atlantic archipelagos, incl. Cape Verde, Canary, Madeira, and Azores islands, formerly or presently part of Portugal or Spain Mad--Madeira, Portuguese islands in Atlantic Ocean north of the Canary Islands Mal--Malaya, peninsular portion of Malaysia south of Thailand, here considered the westernmost part of Malesia Man--Manitoba, Canadian province east of Saskatchewan, west of Ontario and Hudson's Bay, north of North Dakota and Minnesota, and south of District of Keewatin Manc-Manchuria, region of China north of Korea, east of Mongolia, and east and south of the Russian Far East Mar--Mariana Islands, Pacific oceanic islands between Bonins to north and Carolines to south, part of Micronesia Marq--Marquesas Islands, volcanic oceanic islands east of Line and north of Tuamotu islands, part of Polynesia Masc--Mascarene Islands in the Indian Ocean, composed mainly of Mauritius and the French island Reunion Mass--Massachusetts, NE USA Maur--Island of Mauritius in Indian Ocean, which with Reunion and smaller islets constitute the Mascarene Islands Med--Mediterranean Sea area of southernmost Europe and northern Africa north of the Sahara Desert Mex--Mexico, here treated as northernmost republic of Central America, though geologically part of North America Mi--Michigan, NC USA Mic--Micronesia, oceanic islands in Pacific Basin west of Polynesia Mis--Mississippi, SC USA Min--Minnesota, NC USA Mol--Moluccas or Spice Islands, part of Indonesia between Borneo and New Guinea, geologically unstable region between Asian and Australasian continental masses Mong--Mongolia, republic of central Asia north and west of China and south of Siberia Moz-Mozambique, nation on the Indian Ocean north of the Republic of South Africa and east of Zimbabwe nAfr--Northern tier of African republics south of the Mediterranean Sea, including the Atlas Mountains and much of the Sahara Desert, with an essentially Mediterranean flora, here treated botanically as part of Europe Nam-Namibia, a new republic, formerly the trustee territory of South-West Africa governed by the Republic of South Africa, west of Botswana and north of the Republic of South Africa NB--New Brunswick, Canadian Province ne of Maine and w of Nova Scotia and Prince Edward I NCal--New Caledonia NEng--New England Nev--Nevada, W USA

256

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Table V
Continued Nf--Newfoundland, large Canadian maritime island between Gulf of St. Lawrence and Atlantic Ocean, E of Quebec and SE of Labrador, both Labrador and Newfoundland forming Province of Newfoundland NGu--Island of New Guinea with small satellite islands NHeb--New Hebrides, an archipelago between the Solomon and Fiji archipelagos and north of New Caledonia, now the nation of Vanuatu NS--Nova Scotia, Canadian Province E of Maine and New Brunswick NSW--New South Wales, Australian state south of Queensland and north of Victoria Ont--Ontario, central Canadian province between Hudson's Bay and Great Lakes, west of Quebec and east of Manitoba Qld-Queensland, northeasternmost state of Australia with tropical rainforest west of the Great Barrier Reef, especially north of TownsviUe to the Cape York Peninsula Pan-Panama, southernmost republic of Central America between Costa Rica and Colombia Par--Paraguay, South American nation north of Argentina, south of Bolivia, and west of southern Brazil Pat--Patagonia, an arid area mostly in southern Argentina, just north of Fuegia Pen--Pennsylvania,EC USA Phil--Republic of the Philippines, a Malesian archipelago north of Borneo and Celebes (Sulawesi) Poly-Polynes~a, volcanic oceanic islands in Pacific Basin east of Micronesia, here including all the islands south of the Hawaiian Islands, includingthe Cook, Society, Austral, Marquesas, Tuamotu, and Easter islands, etc. PRico--Puerto Rico, a large Antillean island west of Hispaniola Que-Quebec, large Francophone Canadian province n of New England and New Brunswick, W of Labrador, and E of Hudson's Bay Rev--RevillagigedoIslands, the Mexican islands of Socorro and San Benedicto in Pacific Ocean west of Colima Sak--Sahkalin, elongate, north-south island west of Kurile Islands and Sea of Okhotsk, and east of Tatar Strait, part of Russian Far East Salv--El Salvador, small Central American nation south of Guatemala and Honduras and northwest of Nicaragua Sam--Samoa, Pacific islands east of Wallis and Futuna and north of Tonga SAtl--islands in the South Atlantic Ocean, including Ascension, St. Helena, Tristan da Cunha, and Gough islands, all north of the Antarctic Convergence SCruz-Santa Cruz islands, archipelago lying between Solomon and New Hebrides (Vanuatu) archipelagos Seyc--Seychell Islands SHel- St. Helena, a temperate oceanic, volcanic island in the South Atlantic, with many endemic species Sib-Siberia, vast Asian area of Russian Federated Republic, lying south of Arctic Ocean, east of Ural Mountains, north of Kazakhistan and Mongolia, and west of Russian Far East and Chinese Manchuria Soc- Society Islands, Polynesian islands includingTahiti, lying between the Cook and Tuamotu archipelagos Socot--Socotra, a large continental island, belonging to Yemen, in the Indian Ocean off the Somalian Horn of Africa Sol--Solomon Islands, lying east of the Bismarck Archipelago and northwest of the New Hebrides (Vanuatu) Sul--Sulawesi, Indonesian island east of Borneo, formerly called Celebes Sum-- Sumatra, large island of Indonesia south of the Malay Peninsula and west of Java Sur--Surinam, small country on northeastern Atlantic Coast of South America between French Guiana and Guyana Syr-Syria, a nation in the Near East south of Turkey and east of Lebanon Tas--Tasmania, island state of Australia, south of Victoria and Bass Strait, with much cool temperate rainforest and many subantarctic floristic elements on the high plateaus

CLASSIFICATIONAND GEOGRAPHYOF THE FLOWERINGPLANTS Table V Continued

257

Tex--Texas, southern state north of the Mexican state of Tamaulipas, with, like Florida, a number of tropical floristic elements not found in the rest of the USA Transv-- Transvaal, northernmost province of the Republic of South Africa, south of Zimbabwe and west of Mozambique and Swaziland Urug--Uruguay, small South American democracy between Brazil and Argentina Ven--Venezuela, republic in northern South America south of the Caribbean Sea and between Colombia and Guyana Ver--Vermont, NE USA Wyo-Wyoming Zim-Zimbabwe, formerly Rhodesia, an African nation north of Transvaal, south of Zambia, east of Botswana, and west of Mozambique

mer connections with Africa before the origin of the South Atlantic and with Australasia via Antarctica are certainly also involved in its floristic richness. The relative floristic paucity of the larger continent of Africa must be due to extinctions caused by climatic catastrophes (Raven & Axelrod, 1974) as well as to the earlier tectonic loss of India, Madagascar, and Arabia. The surprising floristic richness of Central America, here including Mexico, and Malesia must be due to their critical intercontinental positions. That of Australia and the large continental islands of Madagascar and New Caledonia must, on the other hand, be due to their present long isolation following Mesozoic separation of Australia and Madagascar from Africa and New Caledonia from Australia. Later separation of Australia from Antarctica and South America came early in the Tertiary. The floristic poverty of oceanic islands surely is due to isolation from large continental masses, small size, and relative recentness of volcanic origin. Reasonably consistent with the relative floristic richness of these regions is the relative degree of familial and subfamilial endemism recorded in Table III. Here, however, the rather isolated continents of Africa and South America far surpass the other areas in endemism. African endemism seems even more striking if we add those taxa endemic to Madagascar and the Indian Ocean islands and four families shared only between Africa and Madagascar--a total of 56 families and additional subfamilies. Even larger figures would ensue for South America if we added those 15 essentially South American families and 21 additional subfamilies that have invaded the West Indies often north to southern Florida and Central America north to Panama, Costa Rica, or even southern Mexico. That total of 69 could be enhanced further by adding the 11 essentially South American families barely represented across the South Atlantic in Africa. One can only conclude that South America and adjacent tropical regions to the north are the richest repository on earth for major angiosperm taxa restricted completely or largely to one continent. Table IV summarizes the world distribution of angiosperm families and subfamilies. The largest number of major angiospermous taxa, 280, belong to the subcosmopolitan, pantropical, and widespread temperate categories. Next are the narrower disjuncts between continents and/or oceanic regions, 260 taxa. Finally those major taxa restricted to one continent number 167 taxa to total 708 subfamilies and undivided families. Table V supplies the abbreviations for the geographic areas listed in the synopsis that follows.

258

THE BOTANICAL REVIEW

In the following synopsis of the Angiospermae, within the two subclasses, the hierarchy consists, in descending order, of superorders (-anae), orders (-ales), suborders (-inae), families (-aceae), subfamilies (-oideae), and, in Asteraceae, tribes (-eae). An example in the Dicotyledoneae (Magnoliidae = Annonidae) is: Superorder: Magnolianae (Annonanae) Order: Magnoliales (Annonales) Suborder: Magnoliineae (Annonineae) Family: Magnoliaceae Subfamily: Magnolioideae (Tribe: Magnolieae). This example is presented to eliminate the need to list in every case the hierarchical ranks preceding the respective taxa.

IV. Classification and Distribution of the Families and Subfamilies of the Class Angiospermae (Magnoliopsida)
SUBCLASS: DICOTYLEDONEAE (MAGNOLIIDAE)

Magnolianae (Annonanae) (C; 466/11,910) Magnoliales (incl. Annonales, Laurales, etc.) (C; 281/8410) Winterineae (C; 8/90) Winteraceae (C; 8/90) TropMont-cool temp Au(nQld-Tas) C(-Mex) G nMa Me(Phil-Sol) P(Fern) S(-Fuegia) Z Illiciineae (C; 3/84) Illiciaceae (C; 1/37) (Illicium) TropMont-temp se, eAs--Born, Phil; seUSA; eMex; Ant Schisandraceae (C; 2/47) (Kadsura, Schisandra) TropMont-temp se, eAsBorn, Phil; seUSA Magnoliineae (Annonineae) (C; 178/3250) Magnoliaceae (C; 12/220) TropMont-temp se, eAs-NGu; Mex-seBraz; Ver & sOnt-WInd Magnolioideae (11/220) TropMont-temp se, eAs--NGu; Mex-seBraz & Peru; Mass & sOnt-WInd Liriodendroideae (1/2) (Liriodendron) Temp eAsia; Ver & Mi-F1 & La Degeneriaceae (C; 1/2) (Degeneria) Trop Fiji Is Himantandraceae (C; 1/2) (Galbulimima) TropMont Mol-Bis Is; neQld Eupomatiaceae (C; 1/2) (Eupomatia) TropMont-temp eAu-NGu Annonaceae (C; 132/2300) Trop-warm temp Af(-CapeP) seAs nAu C(-Mex) I(-Masc) Ma Me(-Fiji) eN(-NY, Ont-Fla, Nebr) P(-Soc, Ga!) S(-Urug) Wind Aristolochiaceae (C; 8/400) Trop-temp Af(-Zim) As At(Mac) Au(neQld) C E G Ma Me(-Fiji) N(-Que, BrC) P(-Sam, Rev) S(--Chile) Wind Asaroideae (3/71) (Asarum, Hexastylis, Saruma) Temp Eurasia; Que & BrCFla& La Aristolochioideae (5/330) Trop--temp Af(-Zim) As At(Mac) Au(neQld) C E G Ma Me(-Fiji) N(-NE, Cal) P(-Sam, Rev) S(-Chile) Wind Myristicaceae (C; 17/300) Trop Af seAs Au(neQld) C(-Mex) Ma Me(-Fiji) P(--Car, Tonga) S(-seBraz, Bol) Wind

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259

CaneUaceae (C; 6/21) Trop Mex, sFla & Wlnd-seBraz; eAf; Ma Austrobaileyineae (C; 1/1) Austrobaileyaceae (C; 1/1) (Austrobaileya) TropMont neQld Laurineae (C; 77/2975) Amborellaceae (C; 1/1) (Amborella) TropMont NCal Trimeniaceae (C; 1/5) (Trimenia, incl. Piptocalyx) TropMont eAu G Me(SulFiji) P(--Sam, Marq) Chloranthaceae (C; 4/70) TropMont-warm temp e, seAs-Fiji, NCal, Marq; Ma; sMex & Ant-Par Monimiaceae (C; 32/335) Trop--warm temp Af seAs eAu(Qld-Tas) C(-Mex) G I(-Masc) Ma Me(-Fiji) P(-Tonga) S(-Chile) Wind Z Hortonioideae (1/3) (Hortonia) Trop Sri Lanka Monimioideae (3/18) (Monimia, Palmeria, Peumus) Trop seAs Au(neQld) I(-Masc) Me(-Bis) S(Chile) Mollinedioideae (19/145) Trop Af(--CapeP) seAs Au(neQld) C G I(Masc) Ma Me(-Fiji) P(-Tonga) S(-Urug) Z Atherospermatoideae (7/16) TropMont-warm temp NGu-Tas; NCal-NZ; Peru--Chile & Pat Siparunoideae (1/150) (Siparuna) Trop Ant & sMex-seBraz & Bol Glossocalycoideae (1/3) (Glossocalyx) Trop wAf Gomortegaceae (C; 1/1) (Gomortega) Temp cChile Calycanthaceae (C; 3/7) Trop-temp eAs; neQld; Cal; eUSA Idiospermoideae (1/1) (Idiospermum) Trop neQld Calycanthoideae (2/6) (Calycanthus, Chimonanthus) Temp eAs; Cal; Pen & Ohio--Ha & Mis Lauraceae (C; 31/2490) Trop--temp Af(-Cape) As At(Mac) Au C E(Med) G H I(-Masc, Seyc) Ma Me(-Fiji) N(-Me, Ont) P(-Poly) S(-Chile) Wind Z Lauroideae (30/2470) Trop--temp Af(--Cape) As At(Mac) Au C E(Med) G H I(-Masc) Ma Me(-Fiji) N(-Me, Ont) P(-Tonga, Cocos) S(--Chile) Wind Z Cassythoideae (1/20) (Cassytha) Trop Af(--CapeP) seAs Au C(-Mex) G H I(-Masc, Seyc) Ma Me(-Fiji) N(Fla) P(-Poly) S(-Braz) Wind Z Hernandiaceae (C; 4/64) Trop Af(-Zim) e, seAs Au(eQld) C(-Mex) G I(-Masc, Seyc) Ma Me(-Fiji) P(-Marq) nS(Sur-Bol) Wind Hemandioideae (2/42) (Hernandia, Illigera)Trop Af(-Ang) e, seAs Au(eQld) C(-Mex) G I(-Masc, Seyc) Ma Me(-Fiji) P(-Bon, Marq) nS(Sur-Ecu) Wind Gyrocarpoideae (1/7) (Gyrocarpus)Trop Af(-Zim) seAs Au(neQld) C(-Mex) G I(--Seyc) Me(-Phil, Fiji) P(--Soc) nS(--Col, Ven) Wind Sparattanthelioideae (1/15) (Sparattanthelium) Trop Mex-Braz & Bol Piperineae (C; 14/2010) Lactoridaceae (C; 1/1) (Lactoris)Cool temp Mas a Tierra of Juan Fernandez Is Saururaceae (C; 5/7) Trop--temp se, eAs--Jap & Phil; NEng & Min-Fla & Tex; Cal & Nev-Tex & Mex Piperaceae (C; 8/2000) Trop--subtr Af(--Cape) As At eAu C(-Mex) G H I(Masc, Seyc) Ma Me(-Fiji) N(sFla) P(-Marq, Fern) S(-Chile) Wind Z Piperoideae (4/1000) Trop-subtr Af(--CapeP) se, eAs Au(neQld) C(-Mex) G I(-masc, Seyc) Ma Me(-Fiji) P(-Marq) S(-Par) Wind Z Peperomioideae (4/1000) Trop--subtr Af(--Cape) se, eAs At(Mac) eAu C(-

260

THE BOTANICAL REVIEW

Mex) G H I(-Masc, Seyc) Ma Me(-Fiji) N(sFla) P(Marq, Fern) S(--Chile) Wind Z CeratophyUales (A; 1/6) Ceratophyllaceae (C; 1/6) (Ceratophyllum) Subcosm Af(--CapeP) As(-Jap) At(Az) n, seAu C E G H Ma Me(-Fiji) N(-Alas) P(Mar, Sam, Gal) S(Pat) Wind Z Nelumbonales (C; 1/2) Nelumbonaceae (C; 1/2) (Nelumbo)Trop--temp se, eAs--Phil, NGu; nAu; Mass & eNebr-Fla & eMex; Ant-nCol Paeoniales (B; 2/34) Paeoniaceae (C; 1/33) (Paeonia)Temp Euras(-nAf); BrC & Wyo-BCal Glaucidiaceae (C; 1/1) (Glaucidium)Temp Jap Berberidales (C; 181/3490) Berberidineae (C; 139/2830) Menispermaceae (C; 65/350) Trop--temp Af(CapeP-Sinai) As(-Kor) nAu C G H I(-Maur) Ma Me(-Fiji) N(--Que) P(-Marq) S(-Arg) Wind Lardizabalaceae (C; 8/37) Trop--warm temp se, eAs; cChile & Pat Decaisneoideae (C; 1/2) (Decaisnea)Warm-temp Chi, Him Lardizabaloideae (C; 7/35) (excl. Sargentodoxa)Subtr-temp Him--Chi, nLaos & Vietnam-Kor; cChile & Pat Sargentodoxaceae (B; 1/1) (Sargentodoxa)Trop--warm temp Chi, seAs Berberidaceae (C; 16/540) TropMont-temp eAf(-Mal) As At(Mad) C E(-nAfr) wMe N(--Que) P(Fem) S(-Fuegia) Nandinoideae (C; 1/1) (Nandina) Temp Chi, Jap Berberidoideae (3/520) (Berberis,Mahonia,Ranzania) TrolaMont--cool temla eAf(-Mal) As At(Mad) C E(-nAfr) wMe(Born, Phil) N(-Va, BrC) P(Fern) S(-Fuegia) Leonticoideae (3/8) (Caulophyllum,Gymnospermium,Leontice)Temp seEeAs; eN Epimedioideae (9/11) Temp Euras(-nAf); BrC--Cal; Que & Man-Fla & Tex Hydrastidaceae (C; 1/1) (Hydrastis) Temp eN (NEng, sOnt & Minn-Georgia & eKan) Ranunculaceae (C; 46/1900) Subcosm Af(--Cape) As At(-Mac) Au C E G H I(-Masc) Ma Me(-Fiji) N(--Green) O P(Bon) S(-Fuegia) Wind Z Isopyroideae (Thalictroideae) TropMont-Arct Af(--CapeP) As C E Me(NGu) N(-Green) S(Andes--Arg) Ranunculoideae (incl. Helleborus,Kingdonia)Subcosm Aft-Cape) As(-Kam) At(-Mac) Au C E G H I(-Masc) Ma Me(-Fiji) N(--Green) O P(Bon) S(Fuegia) Wind Z Circaeasteraceae (C; 1/1) TempMont Chi, Him Papaverineae (C; 42/660) Papaveraceae (C; 42/660) Subtr-Arct Af(--Cape) As At(Mac) sAu C E(-nAfr) H N(-Green) P(Bon) S(Andes-Pat) Wind Platystemonoideae (3/4) Temp BrC & Utah-BCal Papaveroideae (18/185) Subtr-Arct As(-Kam) At(Mac) C E(-nAfr) H N(Green) S(Andes-Pat) Wind Eschscholzioideae (3/12) Temp Wash & Utah-nwMex Pteridophylloideae (1/ 1) (Pteridophyllum)Temp Jap

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261

Hypecoideae (1/ 10) (Hypecoum) Temp Euras(-nAfr) Fumarioideae (16/450) SubtrMont-bor Aft--Cape) As(-Kam) At(Mac) C(nwMex) E(-nAfr) N(-Green) P(Bon) Nymphaeanae (C; 8/70) Nymphaeales (C; 8/70) Cabombaceae (C; 2/8) (Brasenia, Cabomba) Trop--cool temp Af e, sAs(-Jap) eAu C N(-Alas) S(-Pat) Wind Nymphaeaceae (C; 6/62) Subcosm Aft--Cape) As Au C E I(-Maur) Ma Me(Bis) N(-Alas) S(-Arg) Wind Nymphaeoideae (3/55) (Nuphar, Nymphaea, Ondinea) Subcosm Aft-Cape) As(-Kam) Au C E I(-Maur) Ma Me(-Bis) N(-Alas) S(-Arg) Wind Euryaloideae (2/3) (Euryale, Victoria) Trop--warm temp se, eAs; Amaz(Par) Barclayoideae (1/4) (Barclaya) Trop Burma-NGu Rafltesianae (B; 10/68) Rattlesiales (B; 10/68) Hydnoraceae (C; 2/15) (Hydnora, Prosopanche) Trop--temp Eth--CapeP; Ma & Reunion; Pat-sBraz & Peru; CRica Rafflesiaceae (C; 8/53) Trop--temp Aft--Cape) sAs At(Can) swAu C(-Mex) E(nAfr) Ma Me(-NGu) swN S(-Pat) Wind(Jam) Mitrastemonoideae (1/2) (Mitrastemon) TropMont-temp Chi-Jap; SumNGu; Mex-Guat Cytinoideae (2/10) (Bdallophyton,Cytinus)Trop--temp Af(--Cape)swAs(Cauc) At(Can) C(Mex-Salv) E(nAfr) Ma Apodanthoideae (2/26) (Apodanthes, Pilostyles) Trop-tempAf(-Zim) swAs(Syr-Iran) swAu C N(sCal-Tex) S(-Pat) Wind(Jam) Ratttesioideae (3/15) (Rafflesia,Rhizanthes, Sapria) Trop Assam, YunnanSum, Born Caryophyllanae (Chenopodianae, Centrospermae) (C; 563/8600) CaryophyUales (Chenopodiales) (C; 563/8600) Caryophyllineae (C; 70/1750) Caryophyllaceae (incl. Geocarpon)(C; 70/1750) Subcosm Aft--Cape) As(-Kam) Aft-Mac) Au C E(-Ice) G H I(-Masc) Ma Me(-NGu) N(-Green) O P S(Fuegia) Wind Z Alsinoideae Subcosm Aft--Cape) As(-Kam) At(-Mac) Au C E(-Ice) G H I(Masc) Ma Me(-NGu) N(-Green) O P(Gal) S Wind Z Paronychioideae Trop--cool temp Aft--Cape) As At(Mac) Au C E I Ma Me N(-Me) P(Fern) S(-Pat) Wind Z Caryophylloideae Trop--Arct Aft-Cape) As(-Kam) At(Mac) Au C E(-Ice) H Ma Me N(--Green) S(-Fuegia) Portulacineae (C; 122/1955) Portulacaceae (C; 19/500) Subcosm Af(--CapeP) As(-Kam) At Au C E(-Ice) G H I(-Masc) Ma Me(Phil-Fiji) N(--Green) O P(-Marq) S(-Fuegia) Wind Z Hectorellaceae (C; 2/2) (Hectorella, Lyallia) Cool temp sZ; Kerguelen Basellaceae (C; 4/40) Trop--warm temp Af seAs C(-Mex) G Ma Me(NGu) N(Tex) P(Gal) S(-Arg) Wind Didiereaceae (C; 4/11) Trop arid swMa Cactaceae (C; 93/1400) Trop-temp Af As(Ceyl) At(Ber) C I(Masc, Seyc) Ma N P(Gal) S(-Pat) Wind

262

THE BOTANICAL REVIEW

Pereskioideae (2/18) (Maihuenia, Pereskia) Trop--temp Mex & Wind-Pat Opuntioideae (4/250) Trop--temp NEng & BrC-Pat; Bermuda; Wind; Gal Cactoideae (87/1130) Trop-temp Af(-Natal) As(Ceyl) C I(Masc, Seyc) MAN(Minn, Alb) P(Gal) S(-Pat) Wind Phytolaccineae (C; 202/2530) Phytolaccaceae (C; 6/55) Trop-temp Af(-Cape) s, eAs(-Jap) C H Ma Me N(Me, Ont) P(Cocos, Gal) S(-Pat) Wind Phytolaccoideae (incl. Lophiocarpus) (5/50) Trop--warm temp Af(-Cape) s, eAs(-Jap) C H Ma Me N(-Me, Ont) P(Cocos, Gal) S(-Pat) Wind Gisekioideae (1/5) (Gisekia) Trop Af-sAs; Ma & Masc Petiveriaceae (Rivinaceae) (C; 6/40) Trop--subtr Af(--CapeP) seAs Au(eQld) C G Me(NHeb) sN(Fla-Ariz) S(-Arg) Wind Agdestidaceae (C; 1/1) (Agdestis) Trop-warm temp Mex--Guat Barbeuiaceae (C; 1/1) (Barbeuia) Trop Ma Achatocarpaceae (C; 2/10) (Achatocarpus, Phaulothamnus) Trop-subtr sTex-Arg Stegnospermataceae (C; 1/3) (Stegnosperma) Trop--subtr wMex--CRica; Ant Nyctaginaceae (C; 30/290) Trop--temp Af(-Namib) sAs nAu C E(wMed-Egypt) G H I(-Masc, Seyc) Ma Me(-Fiji) N(-BrC, Wis) P(-Marcl) S(-Pat) Wind nZ Aizoaceae (C; 141/2040) Trop--temp Af(-Cape) seAs(-Jap) At(Mac) Au C E(Med) G H I(-Masc) Ma Me(-Fiji) sN P(-Poly) S(-Pat) Wind Z Aizooideae (6/80) Trop--warm temp Af(--Cape) sAs At(Mac) Au E(Med) I(Socot) Aptenioideae (Mesembryanthemoideae) (9/90) Warm temp Af(--Cape) [At(Mac) E(Med)-natur.?] Ruschioideae (incl. Caryotophoroideae, Hymenogynoideae) (120/1800) Trop-warm temp Af(-Cape) Au C N(--Ore) S(--Chile) Z Sesuvioideae (4/20) Trop-temp Af seAs nAu C G H I(Masc) Ma Me(-Fiji) N(-NY, Cal) P(-Poly) S(-Pat) Wind Tetragonioideae (2/50) (Tetragonia, Tribulocarpus) Trop-temp Af(--Cape) se, eAs(-Jap) Au(s) G H P(-Poly) S(Chile) Z Halophytaceae (C; 1/1) (Halophytum) Temp arid Arg MoUuginaceae (B; 13/90) Trop--temp Af(-Cape) As At(-Az) Au C G I(Masc, Seyc) Ma Me(-Fiji) sN(Tex-Cal) P(-Micr, Gal) S(-Arg, Chile) Wind Chenopodiineae (C; 169/2360) Chenopodiaceae (C; 104/1510) Subcosm Af(--Cape) As(-Kam) At(-Az, Ber) Au C E G H I(Socot) Ma Me(-NGu) N(--Green) O P(Bon, Gal) S(-Fuegia) Wind Z Chenopodioideae (ind. Dysphania, Microtea)Subcosm Af(--Cape) As(-Kam) At(-Az, Ber) Au C E(Ice--nAfr) G H I(Socot) Ma Me(-NGu) N(--Green) O P(Bon, Gal) S(-Fuegia) Wind Z Salicornioideae Trop-temp Af(--Cape) eAs(-Jap) At(Bet) sAu C E G I(Socot) Ma N(-Alas) P(Fern) S(-Fuegia) Wind Z Salsoloideae Trop--temp Af As(-Jap) At(Mac) E(-nAfr) Ma Amaranthaceae (C; 65/850) Subcosm Af(--CapeP) As At(Mac) Au C E G H I(Masc) Ma Me(-NGu) N(-Me) P(-Marq) S(-Pat) Wind Amaranthoideae Subcosm Af(--CapeP) As At(Mac) Au C E G H I(-Masc) Ma Me(-NGu) N(-Me) P(-Marq) S(-Pat) Wind

CLASSIFICATION AND GEOGRAPHY OF THE FLOWERING PLANTS

263

Gomphrenoideae Trop--warm temp Af As At Au C E G I(-Masc) Ma Me(NGu) N(-Md) P(Gal) S(-Pat) Wind Theanae (C; 540/13,216) Theales (C; 214/5135) Dilleniineae (C; 9/400) DiUeniaceae (C; 9/400) Trop--warm temp Af seAs Au C(-Mex) G I(Seyc) Ma Me(-Fiji) S(-seBraz, Bol) Wind DiUenioideae (5/280) Trop--temp seAs Au G I(Seyc) Ma Me(-Fiji) Tetraceroideae (4/120) Trop Af seAs neAu C(-Mex) G Ma Me(-Bis) S(seBraz, Bol) Wind Theineae (C; 66/2410) Actinidiaceae (C; 3/350) TropMont-temp se, eAs-Fiji; neQld; Mex-Bol Actinidioideae (1/40) (Actinidia) TropMont-temp eAs(-Amur) Saurauioideae (1/300) (Saurauia) TropMont-subtr se, eAs-Fiji; neQld; sMex-Bol Clematoclethroideae (1/10) (Clematoclethra) Temp Chi Paracryphiaceae (B; 1/1) (Paracryphia)Trop Mont NCal Stachyuraceae (C; 1/10) (Stachyurus) TropMont-temp eAs(-Jap); Bon Theaceae (C; 28/500) TropMont-warm temp AfAs(-Jap) At(Mac) Au(neQld) C(-Mex) H Me(-Fiji) seN P(-Rapa) S(-Arg) Wind Temstroemioideae(11/200) TropMont Afse, eAs(-Jap) At(Mac) Au(neQld) C(-Mex) H Me(-Fiji) P(-Rapa) S(-Arg) Wind Theoideae (16/300) TropMont-warm temp As(-Jap) Au C(-Mex) Me(-Bis) seN(-Va) P(Bon) S(-seBraz) Wind Sladenioideae (1/1) (Sladenia) Trop seas Asteropeiaceae (B; 1/7) (Asteropeia) Trop Ma Tetrameristaceae (B; 2/2) (Pentamerista, Tetramerista) Trop wMe; nSAmer (GuayH) Pellicieraceae (C; 1/1) (Pelliciera)Trop Pacific coasts of CRica-Ecu; Caribbean coasts of Nic--Col Chrysobalanaceae (A; 17/495) Trop--warm temp Af(-Natal) seAs nAu C(Mex) G I(-Masc) Ma Me(-SCruz) seN(-SC) P(-Cocos, Bon) S(-seBraz, Bol) Wind Symplocaceae (C; 1/500) (Symplocos) TropMont-warm temp As(-Jap) Au(eQld) C(-Mex) G Me(-Fiji) seN(-Va) P(Bon, Car) S(-Arg) Wind Caryocaraceae (C; 2/23) (Anthodiscus, Caryocar) Trop CRica & Sur-Par Marcgraviaceae (C; 5/100) Trop Mex, Cocos I., & Ant-seBraz, Bol Oncothecaceae (B; 1/2) (Oncotheca) TropMont NCal Aquifoliaceae (C; 1/400) (Ilex, incl. Nemopanthus) TropMont-bor Af(-Cape) As(-Jap) At(Mac) Au(neQld) C(-Mex) E G H Ma Me(-Fiji) eN(-Nf, Minn) P(-Marq) S(-Arg) Wind Phellinaceae (C; 1/10) (Phelline) Trop NCal Sphenostemonaceae (B; 1/7) (Sphenostemon) TropMont Sul-Bis; neQld; NCal Sarraceniineae (C; 3/15) Sarraceniaceae (C; 3/15) TropMont GuayH(-Ven) (Heliamphora); Temp--bor NAmer (Ore--Cal) (Darlingtonia) Lab & Sask-Fla & Tex, (Sarracenia) Clethrineae (B; 5/80) Pentaphylacaceae (C; 1/2) (Pentaphylax) Trop--subtr sChi-seAs & Sum

264

THE BOTANICAL REVIEW

Clethraceae (C; 1/64) (Clethra) TropMont-temp e, seAs--NGu; Mad; MaineAnt; Mex-Arg Cyrillaceae (C; 3/14) (Cliftonia, Cyrilla, Purdiaea) TropMont-warm temp Virginia-Ant-Braz; sMex-Nic Scytopetalineae (C; 55/695) Ochnaceae (incl. Lophira) (C; 40/600) Trop-subtr Afr(--CapeP)seAs Au(neQld) C(-Mex) I(-Maur) Ma Me(-Fiji) S(-Par) Wind Ochnoideae (33/525) Trop--subtr Af(-CapeP) seAs Au(neQld) C(-Mex) I(Maur) Ma Me(-Fiji) S(-Par) Wind Sauvagesioideae (7/75) Trop Af seAs C(-Mex) Ma Me(Mol-NGu) S(-Par) Wind Quiinaceae (C; 4/50) Trop Jam; Bel-sBraz and Bol Scytopetalaceae (C; 5/20) Trop Af Medusagynaceae (C; l/l) (Medusagyne) Trop Seyc (Mah6) Strasburgeriaceae (B; 1/1) (Strasburgeria) TropMont NCal Ancistrocladaceae (B; 1/20) (Ancistrocladus) Trop Af; Cey & sChi-Me Dioncophyllaceae (B; 3/3) (Dioncophyllum, Habropetalum, Triphyophyllum) Trop wAf Nepenthineae (C; 1/70) Nepenthaceae (C; 1/70) (Nepenthes) Trop sChi-NGu, Car & neQld; Seyc-Ma Hypericineae (C; 50/1065) Bonnetiaceae (C; 3/22) (Archytaea, Bonnetia, Ploiarium) TropMont seAsNGu; Ant-Braz & Peru Clusiaceae (incl. Hypericaceae) (C; 45/1010) Subcosm Aft-Cape) As(-Kam) At(-Az) Au C E G I(-Masc) Ma Me(-Fiji) N(-NI) P(-Marq) S(--Chile) Wind Z Kielmeyeroideae (7/47) Trop Bel-Par; Ant Calophylloideae Trop Af seAs neAu C G I(-Masc, Seyc) Ma Me(-Fiji) P(Marq) S(-seBraz) Wind Clusioideae Trop AfseAs neAu C G Ma Me(-Fiji) N(sFla) P(-Tonga, Cocos) S(-seBraz) Wind Moronoboideae (6/36) Trop Af seAs neAu C(-Bel) G Ma S Wind Hypericoideae (9/540) Subcosm Af(--Cape) As(-Kam) At(Mac) Au C E G I(-Masc) Ma Me(-NGu) N(-Nf) P(Rev, Gal) S(-Chile) Wind Z Elatinaceae (C; 2/35) (Bergia, Elatine) Subcosm Af(-Cape) As(-Jap) At(-Az) Au C E G Ma Me(-Fiji) N(-Nf) S(-Chile) Wind Z Lecythidineae (C; 20/270) Lecythidaceae (C; 20/270) Trop Af seAs nAu C(-Mex) G I(-Masc, Seyc) Ma Me(-Fiji) P(-Soc) S(-Par) Wind(Jam) Planchonioideae (6/55) Trop Af(-Nat) As Au(n) G I(-Seyc) Ma Me(-Fiji) P(-Soc) Foetidoideae (1/5) (Foetidia) Trop Pemba-Ma-Masc Lecythidoideae (10/200) Trop Jam; Bel-Par Napoleonoideae (2/11) (Crateranthus, Napoleonaea) Trop wAf Asteranthoideae (1/1) (Asteranthos) Trop Amaz Ericales (C; 132/2650) Ericaceae (C; 99/2245) TropMont-Arct Af(--Cape) As(-Kam) At(-Az) eAu C E G H I(-Masc) Ma Me(-Fiji) N(--Green) P(-Marq) S(-Fuegia) Wind Z

CLASSIFICATION AND GEOGRAPHY OF THE FLOWERING PLANTS

265

Rhododendroideae (15/700) TropMont-Arct As(-Kam) Au(neQld) C E(Ice) I(-Masc) Me(-Sol) N(--Green) P(Bon) S(-Urug) Wind Ericoideae (17/865) TropMont-temp Af(--Cape) At(Mac) E(Ice-nAfr) I(Masc) Ma Vaccinioideae (incl. Arbuteae) (54/660) TropMont-Arct Af(-Transv) As(Kam) At(Mac) eAu(-Tas) C E(-Ice) G H I Ma Me(-Fiji) N(--Green) P(Marq) S(-Fuegia) Wind Z Pyroloideae (3/10) (Chimaphila, Moneses, Pyrola) SubtrMont-Arct As(Kam) C E(-Ice) N(--Green) S(-Ven) Wind Monotropoideae (10/12) SubtrMont--cool temp As(-Sak) C E Me(Sum) N(Alas) S(-Col) Epacridaceae (excl. Wittsteinia) (C; 30/400) Trop--cool temp seAs Au G H Me(-Fiji) O P(-Marq) S(Fuegia) Z Empetraceae (C; 3/6) (Ceratiola, Corema, Empetrum) Temp--Arct As(-Kam) At(-Az, Tris) E(-Ice) N(--Green) O P(Fern) S(Pat, Fuegia) Fouquieriales (C; 1/11) Fouquieriaceae (C; 1/11) (Fouquieria, incl. Idria) Subtr-warm temp arid Cal to Tex & Mex Styracales (Ebenales) (C; 67/1450) Ebenineae (Sapotineae) (C; 56/1300) Ebenaceae (C; 2/500) (Diospyros, Euclea) Trop--temp Aft-Cape) se, eAs(-Jap) neAu C(-Mex) G H I(-Masc, Seyc) Ma Me(-Fiji) N(--Conn) P(-Micr, Tonga) S(-Urug) Wind Lissocarpaceae (C; 1/2) (Lissocarpa) Trop Guy-Bol Sapotaceae (incl. Sarcosperma) (C; 53/800) Trop--warm temp Aft--Cape) seAs At(Can, Mad) neAu C E(nAfr) G H I(-Masc, Seyc) sN(-Va) P(-Micr, Poly) S(-Chile) Wind Z Styracineae (C; 11/ 150) Styracaceae (C; 11/150) TropMont-temp eAs(-Jap) C E(Med) Me(-Sol) sN(Va) P(Car) S(-Arg) W Primulales (C; 77/2870) Primulineae (C; 58/2095) Theophrastaceae (C; 5/95) Trop--subtr sFla, Mex & Wind-Par Myrsinaceae (C; 33/1000) Trop--warm temp Aft-Cape) se, eAs At(-Az) eAu C(-Mex) G H I(-Masc, Seyc) Ma Me(-Fiji) N(sFla) O P(-Marq) S(-Arg) Wind Z Myrsinoideae (incl. Aegiceras) (32/900) Trop--warm temp Aft--Cape) As(Jap) At(-Mac) eAu C(-Mex) G I(-Masc, Seyc) Me(-Fiji) N(sFla) O P(Marq) S(-Arg) Wind Z Maesoideae (1/100) (Maesa) TropAf(--CapeP) e, seAs(-Jap) Au(neQld) G Ma Me(-Fiji) P(-Tonga) Primulaceae (incl. Coris) (C; 20/1000) Subcosm Af(-Cape) As(-Kam) At(Mac) seAu C E(-Ice) G H I(-Masc) Ma Me(-NHeb) N(--Green) O P(Rapa) S(-Fuegia) Wind Z Plumbaginineae (C; 19/775) Plumbaginaceae (C; 19/775) Subcosm Af(--Cape)As At(Ber) n, eAu C E(-Ice) G H I(Socot) Ma Me(-Fiji) N(-Green) P(-Marq) S(-Fuegia) Wind Plumbaginoideae (4/24) Trop--temp Af(--CapeP) As Au C E G H I(-Masc)

266

THE BOTANICAL REVIEW

Ma Me(-Fiji) N P(-Marq) S(--Chile)Wind Staticoideae(incl.Aegialitis)(15/750) SubcosmAf(--Cape)As(-Kam) At(Mac) Au C E(-Ice) G I Ma Me(-NGu) N(--Green) P(Bon) S(-Fuegia) Wind Polygonales (B; 49/1100) Polygonaceae(C; 49/1100) Subcosm Aft--Cape)As(-Kam) At(Mac) Au C E(Ice) G H I(-Masc) Ma Me(-Fiji) N(--Green)O P(-Poly) S(-Fuegia) Wind Z Eriogonoideae (17/316) Temp Alas-Mex; sSAmer (Chile & Pat) Polygonoideae (24/550) Subcosm Aft--Cape) As(-Kam) At(Mac) Au C E(Ice) G H I(-Masc, Seyc)Ma Me(-Fiji) N(--Green)O P(-Poly) S(-Fuegia) Wind Z Coccoloboideae(8/230) Trop-subtr AfAs At(Ber)Au C(-Mex) G Me(NGuSol) N(FIa) O S(-Pat) Wind Z Celastranae (A; 60/875) Celastrales (B; 60/875) Celastraceae (C; 55/855) Subcosm Af(--Cape) As(-Sak) At(Mac, Ber) Au C E G H I(-Masc) Ma Me(-Fiji) N(--Que) P(-Marq) S(-Fuegia) Wind Celastroideae (25/?) Subcosm Af(--Cape) As(-Sak) At(Mac) Au C E G H I(Masc) Ma Me(-Fiji) N(--Que) P(-Marq) S(-Fuegia) Wind Tripterygioideae (5/34) Trop-temp eAs(-Jap) Au C(-Mex) Ma S(-Arg) Cassinoideae (20/?) Trop--subtr Aft--Cape)seAs At(Ber)Au(neQld) C(-Mex) G H I(-Masc) Ma Me(-Fiji) N(sFla) P(Car) S Wind Hippocrateoideae (4/100) Trop--subtr Af(--CapeP) seAs neAu C(-Mex) G I(-Masc) Ma Me(-Fiji) N(sFla) P(Car) S(-Arg) Wind Siphonodontoideae (1/5) SiphonodonTrop seAsia-Bis; neQld Goupiaceae (B; 1/3) (Goupia) Trop Braz, Guy, Sur Lophopyxidaceae (B; 1/2) (Lophopyxis) Trop Mal-Sol, Palau Stackhousiaceae (B; 3/16) Trop--temp Au Me(Sum-NGu) P(Car) Z Macgregorioideae(1/ 1) (Macgregoria) Temp Au Stackhousioideae (2/15) (Stackhousia, Tripterococcus)Trop--temp Au-Tas; Z; Sum, Phil-NGu, Car Malvanae (C; 769/14,610) Malvales (C; 246/3300) Sterculiineae (C; 151/2120) Sterculiaceae (incl. Craigia, Maxwellia) (C; 60/700) Trop--warm temp Af(CapeP) seAs(--Chi)At(Ber) Au C G I(-Masc) Ma Me(-Fiji) sN(-NC, Cal) P(-Poly) S(-Chile) Wind Sterculioideae (12/?) Trop Af seAs(--Chi)Au C(-Mex) G I(-Masc) Ma Me(Fiji) P(-Micr, Tonga) S(-Braz) Wind Byttnerioideae (48/?) Trop--warm temp Af seAs At(Ber) Au C G I(-Masc) Ma Me(-Fiji) sN(-NC, Cal) P(-Poly) S(--Chile)Wind Huaceae (C; 2/3) (Afrostyrax, Hua) Trop wAf (Ghana-Congo) Elaeocarpaceae(excl.Muntingia) (A; 9/350) Trop-temp se, eAs(-Jap) eAu C(Mex) G H I(-Maur) Ma Me(-Fiji) P(-Poly) S(--Chile)Wind Z Plagiopteraceae (A; 1/1) (Plagiopteron)Trop Burma, Thailand Tiliaceae (incl. Goethalsia, Muntingia) (C; 49/450) Trop--temp Af(--CapeP) As(-Jap) At(Ber) nAu C(-Mex) E G I(-Masc) Ma Me(-Fiji) eN(--Que)P(Poly) S(--Chile)Wind Z Brownlowioideae(12/56) Trop Af seAs nAu C G Ma Me(-Fiji) P(--Soc)S(Braz) Wind

CLASSIFICATION AND GEOGRAPHY OF THE FLOWERING PLANTS

267

Tetralicoideae (1/3) (Tetralix) Trop Cuba; nSAmer Tilioideae (35/390) Trop--temp Af(-CapeP) As(-Jap) At(Ber) nAu C E(nAfr) G I(-Masc) Ma Me(-Fiji) N(--Que) P(-Poly) S(--Chile) Wind Z Neotessmannioideae (1/1) (Neotessmannia)Trop Peru Monotaceae (C; 3/21) Trop Af; Ma; Guy-Ven Pakaramaeoideae (1/1) (Pakaramaea)TropMont GuayH (Guy-Ven) Monotoideae (2/20) (Marquesia, Monotes) Trop Af; Ma Dipterocarpaceae (C; 16/550) Trop seAs--NGu; Seyc Sarcolaenaceae (C; 8/28) Trop eMa Diegodendraceae (Rhopalocarpaceae, Sphaerosepalaceae) (B; 3/15) Trop Ma Rhopalocarpoideae (Sphaerosepaloideae) (C; 2/14) (Dialyceras, Rhopalocarpus) Trop Ma Diegodendroideae (1/1) (Diegodendron)Trop Ma Malvineae (C; 95/1180) Bombacaceae (C; 20/180) Trop AfseAs nwAu C(-Mex) Ma Me(--Sol)P(Cocos) S(-Arg) Wind Malvaceae (C; 75/1000) Subcosm Aft--Cape)As(-Jap) At(Ber) Au C E(-nAfr) G H I(-Masc) Ma Me(-Fiji) N(-Mass) P(-Marq) S(-Pat) Wind Z Urticales (excl. Barbeya) (C; 116/2680) Ulmaceae (C; 15/200) Trop--temp Aft--Cape) As(-Kam) At(Ber) nAu C E G I(-Masc) Ma Me(-Fiji) N(-Nt) P(-Marq) S(-Arg) Wind Celtidoideae Trop--temp Af(--Cape) As(-Jap) At(Ber) nAu C E G I(-Masc) Ma Me(-Fiji) N(-Que) P(-Marq) S(-Arg) Wind Ulmoideae TropMont-temp Euras(-Kam); Nf-Pan Moraceae (C; 53/1400) Trop-temp Af(--CapeP) As(-Jap) n, eAu C seE G H I(Masc, Seyc) Ma Me(-Fiji) N(-Vt) P(-Marq, Gal) S(-Arg) Wind Z Cecropiaceae (C; 6/275) Trop Af seAs C(-Mex) Me(-Sol) P(Cocos) S(-seBraz) Wind Urticaceae (C; 39/800) Subcosm Aft-Cape) As(-Kam) At(Mac, Ber) Au C E(Ice) G H I(-Masc, Seyc) Ma Me(-Fiji) N(--Green) O P(-Marq) S(-Fuegia) Wind Z Cannabaceae (C; 3/3) (Cannabis, Humulus, Humulopsis) Temp Euras(-nAf); NS & Mont-NCar & Cal Rhamnales (C; 48/900) Rhamnaceae (C; 45/850) Subcosm Af(--Cape) As(-Jap) At(Mac) Au C E G H I(-Masc, Seyc) Ma Me(-Fiji) N(-Nf, BrC) O P(-Marq) S(-Fuegia) Wind Z Elaeagnaceae (C; 3/50) (Elaeagnus, Hippophae, Shepherdia) Trop--bor As(Jap) Au(neQld) nE(-Fin) Me(-Bis) N(-Alas) P(Bon) Euphorbiales (C; 359/7730) Euphorbiaceae (C; 304/7030) Subcosm Af(--Cape) As(-Sak) At(Ber) Au C E G H I(-Masc) Ma Me(-Fiji) N(--Que) P(-Marq) S(-Fuegia) Wind Z Phyllanthoideae Trop--temp AfAs(-Jap) At(CVerde) Au C E G H I(-Masc, Seyc) Ma Me(-Fiji) sN(-Pa) P(-Marq) S(-Arg) Wind Oldfieldioideae (incl. Picrodendron)Trop--warm temp Af As Au C(-Mex) Ma Me sN(Cal) S Wind(Bah-Jam) Acalyphoideae Trop-temp Aft-Cape) As(-Jap) At Au C E G I(-Masc, Seyc) Ma Me(-Fiji) N(-Que) P(-Soc) S(-Arg) Wind Crotonoideae Trop-temp Af(-Cape) As At(Ber) Au C E G I(-Masc) Ma Me(-Fiji) N(-ND) P(-Tonga) S(-Chile) Wind Z

268

THE BOTANICAL REVIEW

Euphorbioideae Subcosm Aft-Cape) As(-Sak) At(Mac) Au C E G H I(Masc, Seyc) Ma Me(-Fiji) N(-Oue) P(-Poly) S(-Fuegia) Wind Z Pandoideae (C; 4/28) Trop wAf; seAs-Sol Simmondsiaceae (C; 1/1) (Simmondsia) Warm temp Cal-nwMex Dichapetalaceae (B; 4/200) Trop Af seAs Au(neQld) C(-Mex) Ma Me(-Fiji) P(-Tonga) S(-Braz, Peru) Wind Gonystylaceae (C; 3/23) (Aetoxylon, Amyxa, Gonystylus) Trop seAs-Fiji Thymelaeaceae (C; 47/475) Subcosm Aft-Cape) As(-Kam) At(Mac) Au C E G H I(Socot) Ma Me(-Fiji) N(-Que) P(-Marq) S(-Fuegia) Wind Z Aquilarioideae (5/40) Trop Af seAs Au(neQld) G Ma Me(-NHeb) Gilgiodaphnoideae (1/ 1) (Synandrodaphne) Trop wAf Thymelaeoideae (41/435) Trop-temp Af(-Cape) As(-Sak) At(Mac) Au C E G H I Ma Me(-Fiji) N(-Que) P(-Marq) S(-Fuegia) Wind Z Violanae (C; 725/9580) Violales (C; 288/5350) Cistineae (C; 11/225) Bixaceae (C; 1/4) (Bixa) Trop Ant & Mex-Arg Cochlospermaceae (C; 2/20) (Amoreuxia, Cochlospermum) Trop--warm temp Af seAs nAu C Me(Mal, Sum) swN(Tex-Ariz) S(-Par) Wind Cistaceae (C; 8/200) Subtr-temp Af(Som) As(-sChina) At(Mac) C(Mex-Pan) E(-nAfr) N(-Nf) S(Braz--Chile) Wind Violineae (incl. Caricineae, Salicineae) (C; 144/3185) Violaceae (C; 22/900) Subcosm Af(--Cape) As(-Kam) At(-Az) Au(-Tasm) C E(-Ice) G H Ma Me(-Fiji) N(--Green) P(-Tonga, Sam) S(-Fuegia) Wind Z Violoideae (21/890) Subcosm Af(--Cape)As(-Kam) At(Mac) Au(-Tas) C E(Ice) G H Ma Me(-Fiji) N(-Green) P(-Tonga, Sam) S(-Fuegia) Wind Z Leonioideae (1/6) (Leonia) Trop SAmer(-Braz & Peru) Fusispermoideae (1/3) (Fusispermum) Trop Pan, Col & Peru Flacourtiaceae (incl. Neopringlea, Prockia but excl. Abatieae, Alzatea, Lacistemateae, Paropsiae) (C; 79/880) Trop--warm temp Af(-Cape) se, eAs(Jap) eAu C(-Mex) G H I(-Masc, Seyc) Ma Me(-Fiji) N(seTex) P(-Marq) S(--Chile) Wind Lacistemataceae (C; 2/27) (Lacistema, Lozania) Trop Mex-Arg; Ant Salicaceae (C; 3/530) (Chosenia, Populus, Salix) Trop--Arct Af(--Cape) As(Kam) At(Mac) C E(Ice--nAfr) Ma Me(-Java, Born, Phil) N(--Green) S(Pat, Chile) Wind Dipentodontaceae (C; 1/1) (Dipentodon) Subtr-warm temp nelndia, BurmaswChi Peridiscaceae (B; 2/2) (Peridiscus, Whittonia) Trop Braz, Ven Scyphostegiaceae (C; 1/1) (Scyphostegia)TropMont Born Passifloraceae (incl. Abatieae, Paropsieae) (C; 18/630) Trop--warm temp Af(CapeP) seAs At(Ber) n, eAu G I(Seyc) Ma Me(-Fiji) N(-Pa) P(-Tonga, Gal) S(-Arg) Wind Z Turneraceae (C; 8/120) Trop--warm temp swAfAt(Ber) C(-Mex) I(-Masc) Ma seN(-SC) S(-Arg) Wind Malesherbiaceae (C; 1/35) (Malesherbia) Andean Peru-nChile & wArg Achariaceae (C; 3/3) (Acharia, Ceratiosicyos,Guthriea)Warm temp SAf(CapeP) Caricaceae~(C; 4/55) Trop--subtr Af; Mex--Chile; Jam Tamaricineae (C; 7/190)

CLASSIFICATION AND GEOGRAPHY OF THE FLOWERING PLANTS

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Tamaricaceae (incl. Hololachna, Myrtama) (C; 5/100) Subtr-temp CapePnAf & Mac; Euras (Norway-Jap) Frankeniaceae (C; 2/90) (Frankenia, Hypericopsis) Trop--cool temp Aft-Cape) wAs(--Chi) At(-Mac, SHel) Au C sE(-nAfr) swN(Cal) P(Desv) S(-Pat, Fuegia) Begoniineae (incl. Cucurbitineae) (C; 126/1750) Cucurbitaceae (C; 118/825) Subcosm Af(--CapeP) sw--eAs(-Sak) At(Mac) Au C E G H I(-Masc, Seyc) Ma Me(-Fiji) N(--Que, BrC) P(-Soc) S(--Chile) Wind Z Zanonioideae (18/80) Trop Af seAs AuC(-Mex) Me(-Bis) S(-Braz) Wind Cucurbitoideae (100/745) Subcosm Af(--CapeP) sw-eAs(-Sak) At(Mac) Au C E G H I(-Masc, Seyc) Ma Me(-Fiji) N(-Que, BrC) P(-Soc) S(-Chile) Wind Z Begoniaceae (C; 5/920) Trop--subtr Af(--CapeP) As(-nChi) C(-Mex) H I(-Masc, Seyc) Ma Me(-Fiji) S(-Arg) Wind Datiscaceae (incl. Tetrameleae) (C; 3/4) (Datisca, Octomeles, Tetrameles) Trop-temp Crete-sAs; Sri Lanka & sChi-Sol & neQld; Cal & wNev-BCal Brassicales (Capparales) (C; 436/4230) Resedaceae (C; 6/70) Subtr-temp Af(--CapeP) sw, cAs(nwlndia) At(Mac) C(nMex) E(-nAfr) swN(Cal-Tex) P(Guad) Capparaceae (C; 46/930) Trop--temp Af(-CapeP) As(-sJap) Au C E(Med) G H I(-Masc) Ma Me(-Fiji) N(-BrC) P(-Marq) S(-Chile) Wind Tovarioideae (C; 1/2) (Tovaria) TropMont Mex-Peru & Bol; Ant Pentadiplandroideae (B; 1/2) (Pentadiplandra) Trop wAf Koeberlinioideae (C; 1/1) (Koeberlinia) Trop Bol; warm temp swUSA-Mex Capparoideae (incl. Dipterygium, Oceanopapaver) (C; 30/640) Trop-warm temp Af(-CapeP) As(-sJap) Au C E(Med) G H I(-Masc) Ma Me(-Fiji) sN(Fla) P(-Poly) S(-Pat) Wind Cleomoideae (incl. Buhsia, Podandrogyne) (C; 13/285) Trop--temp Af(CapeP) As Au C E I(-Masc) Ma Me(-Fiji) N(-BrC) P(-Marq) S(--Chile) Wind Brassicaceae (excl. Dipterygium) (C; 376/3200) Subcosm Af(-Cape) As(-Kam) At(Mac) Au C E(-Ice) G H I(-Masc) Ma Me(-Fiji) N(-Green) O P(-Marq) S(-Fuegia) Wind Z Salvadoraceae (A; 3/12) (Azima, Dobera, Salvadora) Trop Afr(-Cape); Ma; se, eAs-Mal & Phil Gyrostemonaceae (A; 5/17) Temp Au Batales (A; 1/2) Bataceae (C; 1/2) (Batis) Trop--warm temp nAu-sNGu; SCar-Wlnd-Braz; Cal-nPeru & Gal Santalanae (B; 162/2040) Santalales (C; 143/1995) Olacaceae (C; 25/250) Trop--warm temp Af seAs(-Jap) Au(n-Tas) C(-Mex) G I(-Masc, Seyc) Ma Me(-Fiji) seN(Ha) P(-Soc) S(-Pat) Wind Olacoideae (23/240) Trop--warm temp Af seAs(-sJap) Au(n-Tas) C(-Mex) G I(-Masc, Seyc) Ma Me(-Fiji) seN(Ha) P(-Soc) S(-Pat) Wind Octoknemoideae (1/6) (Octoknema) Trop Af Erythropaloideae (1/2) (Erythropalum) Trop seAs-Java, Sul Opiliaceae (C; 10/30) Trop Af seAs Au(neQld) C(-Mex) Ma Me(-Bis) S(-Arg)

270

THE BOTANICAL REVIEW

Medusandraceae (C; 1/2) (Medusandra) Trop wAf Santalaceae (ind. Okoubaka) (C; 30/400) Subcosm Aft--Cape)As(-Jap) At(Can) Au C E G H I(Socot) Ma Me(-Fiji) N(-Alas) P(-Marq, Fern) S(-Fuegia) Z Misodendraceae (C; 1/11) (Misodendrum) Cool temp SAmer(-Fuegia) Loranthaceae (C; 65/850) Trop-temp Af As(-Jap) Au C(-Mex) sE(-nAfr) G I(-Masc, Seyc) Ma Me(-Fiji) P(-Marq) S(-Pat, Chile) Wind Z Eremolepidaceae (C; 3/12) (Antidaphne [incl. Eremolepis], Eubrachion, Lepidoceras) TropMont-warm temp Ant & Mex-Arg Viscaceae (C; 8/440) Subcosm Af(--Cape) As(-Jap) At(Mac) Au C E G H I(Masc, Seyc) Ma Me(-Fiji) N(-Alas, NO P(-Marq) S(sBraz, Par) Wind Z Balanophorales (B; 19/45) Balanophoraceae (C; 18/43) Trop--warm temp seAf seAs(--Chi)neAu C(-Mex) G Ma Me(-Fiji) P(-Rapa) S(-seBraz) Wind Z Mystropetaloideae (1/1) (Mystropetalon) Trop--warm temp Af(CProv) Dactylanthoideae (2/2) (Dactylanthus, Hachettea) Trop NCal; subtr Z Sarcophytoideae (2/3) (Chlamydophytum, Sarcophyte) Trop--warm temp Af(--CProv) Helosidoideae (6/12) Trop seAs--Mol; Ma; Mex-se Braz; Ant Lophophytoideae (4/8) Trop--subtr SAmer(--Chile);Gal Balanophoroideae (3/17) (Balanophora, Langsdorffta, Thonningia) Trop Af seAs(--Chi) Au(neQld) C(-Mex) G Ma Me(-Fiji) P(-Rapa) S(-seBraz) Cynomoriaceae (B; 1/1-2) (Cynomorium) Warm temp Can-Egy; Somalia & Palestine--Mong, Chi Geranianae (B; 189/5560) Linales (C; 56/865) Humiriaceae (C; 8/50) Trop wAf; Nic, CRica & Cocos I-seBraz & Peru Ctenolophonaceae (C; 1/3) (Ctenolophon) Trop wAf; seAs--NGu Hugoniaceae (C; 6/55) Trop Af-Ma & Masc; seAs--Fiji & NCal Ixonanthaceae (B; 4/33) Trop Af; sChi-NGu; Guy-Braz Linaceae (C; 6/210) Subcosm Aft--Cape) As(--Sib) At(Mac) Au C E(-Ice) Ma N(--Green) P(Gal) S(--Chile)Wind Z Erythroxylaceae (C; 2/250) (Erythroxylum, Nectaropetalum) Trop Af(-CapeP) seAs nAu C(-Mex) G I(-Masc, Seyc) Ma Me(--Sol) S(-Arg) Wind Zygophyllaceae (C; 28/255) Trop--warm temp Aft--Cape) As(-Jap) At(Mac) Au C E(-nAfr) G H I(-Masc) Ma Me(-NGu) sN(-Utah) P(-Marcl) S(-Pat) Wind Peganoideae (1/6) (Peganum) Temp Med-Mong; Tex-Mex Morkillioideae (Chitonioideae) (3/4) (Morkillia, Viscainoa, Sericodes) Warm temp Mex Tetradiclidoideae (1/1) (Tetradiclis) Temp arid Egy-sw & cAs Tribuloideae (4/60) Trop--temp Af(-Cape) As(-Jap) At Au C E(Med) G H I(-Masc, Seyc) Ma Me(-NGu) N(Ill--Cal) P(-Marq) S(-Arg) Wind Neoluederitzioideae (2/2) (Neoluederitzia, Sisyndite) Warm temp NamCapeP Zygophylloideae (15/175) Trop-temp Aft--Cape) As At(Mac) Au C E(Med) sN(Fla--Cal) S(-Pat) Wind Augeoideae (1/1) (Augea) Subtrop--warm temp SAf (CapeP, Nam) Nitrarioideae (1/8) (Nitraria) Temp nAf & seEur-Sib; sAu Balanitaceae (B; 1/9) (Balanites) Trop--warm temp Natal-nAf-Burma

CLASSIFICATION AND GEOGRAPHY OF THE FLOWERING PLANTS

271

Rhizophorales (B; 12/84) Rhizophoraceae (C; 12/84) (excl. AnisophyUeeae)Trop-subtr Af(-CapeP) se, eAs(-sJap) At(Ber) nAu C(-Mex) G I(-Masc, Seyc) Ma Me(-Fiji) seN(Fla) P(-Gal, Marq) S(-seBraz, nePeru) Wind Geraniales (B; 29/2365) Oxalidaceae (incl. Averrhoeaceae, Lepidobotrys) (C; 6/890) Trop-cool temp Af(-Cape) As(-Jap) C E(-Ice) I(-Masc) Ma Me(-Java, Cel) N(-Que, Wash) O P(Gal) S(-Fuegia) Wind Geraniaceae (B; 14/775) Subcosm Af(-Cape) As(-Kam) At(-Mac, s) Au C E(Ice) H I(-Masc) Ma Me(-NGu) N(-Green) O S(-Fuegia) Z Geranioideae (incl. Hypseocharis) (6/750) Suhcosm Af(-Cape) As(-Kam) At(-Mac, s) Au(-Tas) C E(Ice-nAfr) H I(-Masc) Ma Me(-NGu) N(Green) O S(Andes-Fuegia) Z Biebersteinioideae (1/5) (Biebersteinia) Temp Greece-wChi Dirachmoideae (1/1) (Dirachma) Arid Socotra Vivianioideae (4/6) Subtr-temp SAmer(-Pa0 Ledocarpoideae (2/11) (Balbisia, Wendtia) Subtr-Andean sSAmer Balsaminaceae (C; 5/600) Trop-bor Af(-Cape) As(-Kam) C(CRica) E I(Seyc) Ma Me(-Sol) N(-Alas) Tropaeolaceae (C; 3/89) (Magallana, Tropaeolum, Trophaeastrum) TropMont Mex-cool temp Chile(-Fuegia) Limnanthaceae (C; 1/11) (Limnanthes, incl. Floerkea) Temp NAmer (Que & BCol-Tenn & esp. Cal) Polygalales (Malpighiales) (C; 92/2245) Malpighiaceae (C; 66/1200) Trop-subtr Af(-CapeP) seAs Au(neQld) C G Ma Me(-Fiji) sN P(Car) S(-Arg) Wind Byrsonimoideae Trop--subtr Mex-Par & Bol; sFla-Wlnd Gaudichaudioideae Trop Af(-CapeP) seAs Au(neQld) C(-Mex) G Ma Me(Fiji) P(Car) S(-Arg) Wind Malpighioideae Trop-subtr Tex-Ariz; Mex-Peru; Wind Trigoniaceae(C; 3/26) (Humbertodendron, Trigonia, Trigoniastrum) Trop seAsSum, Born; Ma; Mex-seBraz & Par Vochysiaceae (C; 7/200) Trop wAf; sMex-seBraz & Par Polygalaceae (incl. Diclidanthera, Xanthophyllum) (C; 15/800) Subcosm Af(Cape) As(-Man) At(-Az) Au(-Tas) C E I Ma Me(Sol) N(-Green) P(Mar, Gal) S(-Fuegia) Wind Krameriaceae (B; 1/17) (Krameria) Trop-warm temp Tex & Cal-Arg & Chile; Ga & Fla-Wlnd Rutanae (C; 1129/24,300) Rutales (Sapindales) (C; 1129/24,300) Rutineae (C; 326/3545) Rutaceae (C; 154/925) Subcosm Af(--Cape)As(-Man) At(Mac) Au C sE(-nAfr) G H I(-Masc) Ma Me(-Fiji) N(-Que) P(-Soc) S(-Chile) Wind Z Rutoideae (incl. Toddalioideae ) (117/?) Subcosm Af(-Cape) As(-Man) At(Mac) Au C sE(-nAf) G H I(-Masc) Ma Me(-Fiji) N(-Que) P(-Soc) S(-Chile) Wind Z Citroideae (=-Aurantioideae) (30/?) Trop-warm temp Af(-CapeP) e, seAs(Jap) eAu C(-Mex) G Ma Me(Sol) P(Mar) Flindersioideae (2/17) (Chloroxylon, Flindersia) Trop seAs-NGu; eAu; NCal

272

T H E B O T A N I C A L REVIEW

Spathelioideae (4/23) (incl. Harrisonia) Trop Bahamas-Amaz Dictyolomatoideae (1/2) (Dictyoloma) Trop Braz-Bol Rhabdodendraceae (B; 1/4) (Rhabdodendron) Trop Amaz Cneoraceae (C; 1/3) (Cneorum) Temp Med & Can; trop eCuba Simaroubaceae (C; 27/192) Trop--warm temp Af se, eAs(-Jap) Au(eQld) C(Mex) G I(--Seyc) Ma Me(-Fiji) sN(Fla, Ariz-Cal) P(-Micr, Gal) S(-Arg) Wind Simarouboideae (excl. Harrisonia) (20/116) Trop--warm temp Af e, seAs(Jap) Au(eQld) C G I(Seyc, Masc) Ma Me(-Fiji) sN(Fla, Ariz--Cal) P(wcPoly) S(-Arg) Wind Kirkioideae (1/8) (Kirkia) Trop & sAf(-Transv) Irvingioideae (incl. Allantospermum) (4/23) Trop Af; Ma; seAs-Born Picramnioideae (1/40) (Picramnia) Trop-subtr Mex-Par; sFla-Wlnd Alvaradoideae (1/5) (Alvaradoa) Trop--subtr Mex-Arg; sFla-Wlnd Ptaeroxylaceae (C; 2/5) (Cedrelopsis, Ptaeroxylon) Trop SAf (CapeP); Ma Meliaceae (C; 52/1310) Trop--warm temp Af(--CapeP) se, eAs(-Jap) neAu C(Mex) G I(-Masc) Ma Me(-Fiji) seN(sFla) P(--Cook) S(-Par) Wind Z Melioideae (incl. Nymania) (37/1260) Trop--warm temp Af(--CapeP)e, seAs(Jap) neAu C(-Mex) G I(-Masc) Ma Me(-Fiji) P(--Cook) S(-Par) Wind Z Quivisianthoideae (1/1) (Quivisiantha) Trop Ma Capuronianthoideae (1/1) (Capuronianthus) Trop Ma Swietenioideae (13/47) Trop-warm temp Afe, seAs neAu C(-Mex) Ma Me(Fiji) seN(sFla) P(-Tonga) S(-Par) Wind Burseraceae (C; 17/500) Trop-warm temp Af(-CapeP) seAs Au(neQld) C(Mex) G I(-Maur) Ma Me(-Fiji) sN(Fla, Ariz-Cal) P(-Tonga, Gal) S(-Par, sBraz) Wind Anacardiaceae (incl. Blepharocarya, Dobinea, Julianiaceae) (C; 70/600) Tropcool temp Aft-Cape) As(-Sak) At(Mac) nAu C sE(-nAfr) G H I(-Masc, Seyc) Ma Me(-Fiji) N(-Que, BrC) P(-Soc) S(--Chile) Wind Z Leitneriaceae (B; 1/1) (Leitneria) Warm temp seUSA (Missouri-nFla and eTex) Tepuianthaceae (A; 1/5) (Tepuianthus) TropMont Ven-Col (GuayH) Coriariineae (C; 1/5) Coriariaceae (C; 1/5) (Coriaria)TropMont-temp se, eAs(-Jap) C(-Mex) E(Med) Me(-Fiji) P(-Soc) S(Andes-Chile) Z Sapindineae (C; 155/2420) Sapindaceae (incl. Filicium) (C; 147/2215) Trop-warm temp Af(-Cape) seAs(Jap) At(Ber) Au C G H I(-Masc) Ma Me(-Fiji) sN P(-Marq, Gal) S(Chile, Arg) Wind Z Dodonaeoideae (C; 1/50) (Dodonaea) Trop--warm temp Af seAs At(Ber) Au C H I(Masc) Ma Me(-Fiji) sN(Fla-Ariz) P(Marq, Gal) S Wind Z Koelreuterioideae (28/70) Trop-warm temp Aft-Cape) eAs Au C I(-Masc) Ma Me(-Fiji) sN S(--Chile) Wind Z Stylobasioideae (1/2) (Stylobasium) Temp WAu Emblingioideae (1/1) (Emblingia) Temp WAu Sapindoideae (112/1880) Trop-warm temp Af(-CapeP) e, seAs(-Jap) At(Ber) Au C G H I(-Masc) Ma Me(-Fiji) sN(--Ga, Karts) P(-Marq, Gal) Hippocastanoideae (C; 2/15) (Aesculus, Billia) Trop-temp seEur--eAs(-Jap); Penn & Mi-Fla & Tex; Cal-Ecu

CLASSIFICATION AND GEOGRAPHY OF THE FLOWERING PLANTS

273

Aceroideae (C; 2/200) (Acer, Dipteronia)Trop-temp Euras(-nAf)-Java, Sul; Alas-Guat Sabiaceae (C; 3/160) Trop-temp e, seAs-Sol; Mex-Peru; Wind; Rev Meliosmoideae (2/105) (Meliosma, Ophiocaryon)Trop-temp As(-Jap)-NGu; Mex-Peru; Wind; Rev Sabioideae (B; 1/55) (Sabia) Trop-temp EAs(-Jap)-Sol Melianthaceae (C; 2/15) (Bersama, Melianthus) Trop Afto temp Cape Akaniaceae (C; 1/1) (Akania) Subtrop eAu(NSW--Qld) Bretschneideraceae (C; 1/2) (Bretschneidera) Temp Chi (incl. Taiwan) Moringaceae (B; 1/14) (Moringa) Trop--subtr Nam; Ma; NAf& Sore-India & sChi Fabineae (C; 647/18,325) Surianaceae (A; 1/1) (Suriana) Trop-subtr Coastal Af(-Moz) seAs(-sChina) At(Ber) Au(neQld) C(-Mex) G I(-Masc, Seyc) Ma Me(-NGu) N(sFla) P(Poly) nS(-eBraz) Wind Connaraceae (C; 16/325) Trop Af(-Natal) seAs Au(neQld) C(-Mex) I(-Masc) Ma Me(-Fiji) P(-Tonga) S(-seBraz, Bol) Wind Connaroideae (15/322) Trop Af(-Natal) seAs Au(neQld) C(-Mex) I(-Masc) Ma Me(-Fiji) P(-Tonga) S(-seBraz, Bol) Wind Jollydoroideae (1/3) (Jollydora) Trop wAf Fabaceae (C; 630/18,000) Subcosm Aft--Cape)As(-Kam) At(Mac, Bet) Au C E(-Ice) G H I(-Masc) Ma Me(-Fiji) N(-Green) P(-Marq) S(-Fuegia)Wind Z Caesalpinoideae (150/2700) Trop-warm temp Af(--CapeP) se, eAs(-Jap) At(Ber) neAu C sE G H I(-Masc, Seyc) Ma Me(-Fiji) N(-Vt, Minn) P(Marq) S(-Pat) Wind Mimosoideae (40/2500) Trop-warm temp Af(-Cape) As(-Jap) At(Ber) Au C G H I(-Masc, Seyc) Ma Me(-Fiji) N(-Va, ND) P(-Tonga) S(-Pat) Wind Swartzioideae (11/185) Trop Africa; Ma; sMex-Braz, Bol; Wind Faboideae (429/12,615) Subcosm Af(-Cape) As(-Kam) At(-Mac, sAtl) Au C E(-Ice) G H I(-Masc, Seyc) Ma Me(-Fiji) N(--Green) P(-Marq) S(Fuegia) Wind Z Proteanae (C; 75/1050) Proteales (C; 75/1050) Proteaceae (C; 75/1050) Trop-temp Af(-Cape) seAs(-Jap) Au(-Tas) C(-Mex) G Ma Me(-Fiji) P(Car) S(-Pat) Z Persoonioideae (7/43) Trop--temp Au(-Tas); NCal; Z Proteoideae (26/?) Trop-warm temp Af(-Cape) Au Ma Sphalmioideae (1/1) (Sphalmium) Trop neQld Camarvonioideae (1/2) (Carnarvonia) Trop Qld GreviUeoideae (40/?) Trop-warm temp Af(Cape) se, eAs(-sJap) Au C(-Mex) G Ma Me(-Fiji) P(-Sam) S(-Pat) Z Rosanae (B; 381/6720) Hamamelidales (C; 35/136) Trochodendrineae (C; 4/6) Trochodendraceae (C; 2/2) Subtr-temp eAs Trochodendroideae (C; 1/1) (Trochodendron) Temp Jap, Kor, Taiwan Tetracentroideae (C; 1/1) (Tetracentron) SubtrMont-temp Nepal & Burmasw and cChi

274

THE BOTANICAL REVIEW

Eupteleaceae (C; 1/2) (Euptelea) SubtrMont-temp Assam--cChi& Jap Cercidiphyllaceae(C; 1/2) (Cercidiphyllum) Temp wChi, Kor, & Jap Hamamelidineae (C; 31/ 130) Platanaceae (C; 1/9) (Platanus) Trop--temp Balkans; Him; Laos; Maine & Minn-Fla & Tex-Mex Hamamelidaceae (C; 30/120) Trop--temp eAf(--CapeP) sw, e, seAs(-Jap) Au(neQld) C Ma Me(-Bis) eN P(Bon) S(Col) Hamamelidoideae (22/100) TropMont-temp eAf(--CapeP)w, se, eAs(-Jap) Au(neQld) C Ma Me(-Bis) eN(--Que)P(Bon) S(Col) Rhodoleioideae (1/1) (Rhodoleia) Trop--subtr Yunnan & Hong Kong-Sum Exbucklandioideae (4/5) (incl. Chunia, Disanthus, Mytilaria) Trop--temp e, seAs(-Jap)-Sum Altingioideae (=Liquidambaroideae) (3/12) (Altingia, Liquidambar, Semiliquidambar) TropMont-temp sw & eAs(-Taiwan)-Java; Conn-Hond Casuarinales (C; 4/70) Casuarinaceae(C; 4/70) Trop-temp seas Au(-Tasm) G I(-Masc, Seyc)Ma(nat?) Me(Mal-Phil, Fiji) P(-Marc0 Balanopales (Buxales) (C; 8/120) Buxineae (B; 6/105) Buxaceae (excl. Simmondsia) (C; 5/103) Trop-temp Af(-CapeP) As(-Jap) C(Mex) E I(Socot) Ma Me(Java, Born, Phil) seN wS(-Bol) Wind Buxoideae (4/100) Trop-temp Af(-CapeP) As(-Jap) C(-Mex) sE I(Soco0 Ma Me(Java, Born, Phil) seN(Ky-Fla) nwS Wind Styloceratoideae (1/3) (Styloceras) Trop Andes (Col-Bol) Didymelaceae (B; 1/2) (Didymeles) Trop Ma Balanopineae (Daphniphyllineae) (C; 2/17) Daphniphyllaceae (C; 1/9) (Daphniphyllum) Trop--temp e & seAs(-Jap)-Bis Balanopaceae (C; 1/8) (Balanops) Trop neQld; NHeb-Fiji; NCal Bruniales (C; 17/115) Roridulaceae (C; 1/2) (Roridula) Temp SAf(CapeP) Bruniaceae (C; 12/75) Temp SAf(Cape-Natal) Geissolomataceae (C; 1/1) (Geissoloma) Temp SAf(swCapeP) Grubbiaceae (C; 1/3) (Grubbia) Temp SAf(CapeP) Myrothamnaceae (C; 1/2) (Myrothamnus) Trop-subtr Af(Angola & TransvNam & Natal); Ma Hydrostachyaceae (C; 1/30) (Hydrostachys) Trop Af(Zaire & Tanz)-SAf(-Natal); Ma Juglandales (A; 12/100) Juglandineae (C; 9/60) Rhoipteleaceae (C; 1/1) (Rhoiptelea) Trop Chi, Vietnam Juglandaceae (C; 8/59) Trop--temp seEurlJap; seAs-NGu; Que & NDak-Andean SAmer(-Arg) & eBraz; Wind Myricineae (C; 3/40) Myricaceae(incl. Canacomyrica) (C; 3/40) Trop--bor Aft-Cape) se, eAs(-Kam) At(Mac, Ber) C wE G Ma Me(Born-NGu) N(-Alas) S(--Chile)Wind Betulales (Fagales) (C; 17/945) Ticodendraceae (A; 1/1) (Ticodendron) TropMont CAmer(Guat-Pan) Betulaceae (C; 6/157) Temp-Arct Euras(Ice-nAf-eAs); Green-Arg

CLASSIFICATION AND GEOGRAPHY OF THE FLOWERING PLANTS

275

Betuloideae (2/95) (Alnus, Betula) TropMont-Arct Euras(Ice-nAfr-KamseAs); Green-Arg Coryloideae (4/62) Temp--bor Euras(-Jap); NAmer--CAmer Nothofagaceae (C; 1/35) (Nothofagus) TropMont--cool temp eAu(-Tasm) G Me(NGu-Bis) sS(Chile, Arg) Z Fagaceae (C; 9/700-800) TropMont-temp Euras(nAf--Sak)-Bis; Que & Vancouver I--Col & Wind Castanoideae (4/300) TropMont-temp Euras(nAf-Jap)-Bis; Maine & WashFla & Tex Fagoideae (5/400) TropMont-temp Euras(nAfr--Sak)--NGu; Que & Vancouver I--Col & Wind Rosales (C; 110/2050) Rosaceae (C; 100/2000) Subcosm Aft-Cape) As(-Kam) At(-Az) Au C E(Ice-nAfr) G H I(-Masc) Ma Me(-Fiji) N(--Green) O P(-Poly) S(-Fuegia) Wind Z Spiraeoideae (incl. Lindleya, Lyonothamnus, Vauquelinia)Temp-Arct Euras(-Kam); Alas-Bol Quillajoideae (2/8) (Kageneckia, Quillaja) SubtrMont-temp Peru-Chile Rosoideae Subcosm Af(--Cape) As(-Kam) At(-Trist, Mac) Au C E(-Ice) G H I(-Masc) Ma Me(-Fiji) N(--Green) O P(Fern) S(-Fuegia) Wind Z Amygdaloideae (Prunoideae) (incl. Exochorda) TropMont-bor Af As(-Sak) At(Mac) Au(neQld) C E Ma Me(-Sol) N(-Lab, BrC) Maloideae (28/1110) TropMont-bor As(-Kam) At(Mad) C E(-nAfr) H N(Alas) P(-Poly) S(Andes-Bol) P(Rev) S(-Urug) Wind Neuradaceae (B; 3/10) (Grielum, Neurada, Neuradopsis) Subtr-temp Af(NamCape); nAfr & Arab-India & Afgh Crossosomataceae (Apacheria, Crossosoma, Glossopetalon)(C; 3/7 or 8) Temp Wash & Idaho-Mex (incl. Guad I) AnisophyUeaceae (A; 4/34) Trop Af; Ma; seAs-Sum & Born; Amaz Saxifragales (C; 92/2565) Tetracarpaeaceae (B; 1/1) (Tetracarpaea) Cool temp Tas Crassulaceae (C; 35/1500) Subcosm AfAs(-Kam) At(Mac) sAu C E(Ice-nAfr) I(-Masc) Ma N(-Green) O P(Bon) S(-Fuegia) Wind Z Sedoideae (incl. Semperviveae, Echeverieae) TropMont-Arct Af As(-Kam) At(Mac) C E(Ice-nAfr) N(-Green) P(Bon) S(-Bol) Cotyledonoideae (incl. Kalanchoeae) Trop-temp Af(-CapeP) swAs At(Mac) E(-nAfr) I(-Masc) Ma Crassuloideae Subcosm Af(-Cape) As(-Jap) At(Mac) sAu C E(Ice-nAfr) Ma N(-Alas) O S(-Fuegia) Wind Z Cephalotaceae (C; l/l) (Cephalotus) Temp swWAu Penthomceae (C; 1/2) (Penthorum)Subtr-temp Indochina-eAs(-Manc); Maine & Minn-Fla & Tex Saxifragaceae (C; 30/550) TropMont-Arct As(-Kam) At(Mad) C E(-Ice) N(Green) S(Andes--Fuegia) Astilboideae (3/40) (Astilbe,Astilboides, Rodgersia) Temp eAs(-Amur); Virginia & Kentucky to Georgia Saxifragoideae (27/510) TropMont-Arct As(-Kam) At(Mad) C E(-Ice) N(Green) S(Andes-Fuegia) Francoaceae (C; 2/2) (Francoa, Tetilla) Temp Chile

276

T H E B O T A N I C A L REVIEW

Grossulariaceae(C; 1/150) (Ribes)Temp-bor Euras(Ice-nAfr-Sak);Alas-Andes & Fuegia Vahliaceae (C; 1/5) (Bistella) Trop--warm temp Af(-CapeP) swAs(-nwIndia) E(nAfr) Eremosynaceae (C; 1/1) (Eremosyne) Temp swW Au Lepuropetalaceae (C; 1/1) (Lepuropetalon)Subtr-temp NCar--Georgia& Tex; Mex; Urug--Chile Parnassiaceae (B; 1/50) (Parnassia) Temp-Arct Euras(Ice-nAfr-Jap); GreenMex Stylidiaceae (B; 6/170) Temp-subAntarct seAs(-sChi) Au(-Tas) Me(-NGu) O S(Fuegia) Z Donatioideae (1/2) (Donatia) Cool temp-subAntarct Tas; Z; Fuegia Stylidioideae (5/166) Trop-subAntarct seAs(-sChi) Au(-Tas) Me(-NGu) O S(Fuegia) Z Droseraceae (C; 4/110) Subcosm Af(-Cape) As(-Kam) Au C E(-Ice) G H Ma Me(-NGu) N(-Green) O P(Car) S(-Fuegia) Wind Z Greyiaceae (C; 1/3) (Greyia) Subtr-warm temp SAf(Transvaal-CapeP) Diapensiaceae (excl. Diplarche) (B; 6/20) Temp--Arct-alp Euras(Ice-KamHim); Green--Georgia Podostemales (B; 50/140) Podostemaceae (B; 50/140) Trop-temp Af(-Natal) se, eAs(-Jap) Au(neQld) C(-Mex) I(-Masc) Ma Me(-NGu) eN(-Que) S(-Urug) Wind Tristichoideae(5/10) Trop Af(Transv-Egy)seAs C(-Mex) I(-Maur) Ma Me(Phil) S(-Urug) Wind Podostemoideae (45/130) Trop-temp Af(-Natal) se, eAs(-Jap) Au(neQld) C(-Mex) I(Masc) Ma Me(-NGu) eN(Que-La) S(-Urug) Wind Cunoniales (C; 33/470) Cunoniaceae (C; 27/410) TropMont-temp Af(Cape) seAs e, swAu(-Tasm) C(sMex) G I(-Masc) Ma Me(-Fiji) P(-Marq) S(-Fuegia) Wind Z Cunonioideae (C; 24/350) Trop Mont-temp Af(Cape) seAs e, swAu(-Tasm) C(-sMex) G I(-Masc) Ma Me(-Fiji) P(-Marq) S(-Fuegia) Wind Z Baueroideae (C; 1/3) (Bauera) Temp eAu(-Tas) Eucryphioideae (C; 1/6) (Eucryphia) Subtr-temp Au(Qld-Tas); Chile & Arg Brunellioideae (C; 1/52) (Brunellia) TropMont sMex-Andean SAmer(Bol); Ant Davidsoniaceae (C; 1/1) (Davidsonia) Subtrop Qld, NSW Staphyleaceae (C; 5/60) TropMont-temp Euras(-Manc)-Bis; Que & MinnPeru; Wind Staphyleoideae (3/55) (Euscaphis, Staphylea, Turpinia) TropMont-temp Euras(-Manc)-Bis; Que & Minn-Peru; Ant Tapiscioideae (2/5) (Huertea, Tapiscia) Trop-temp se, eAs(-Chi); Ant-Ven Cornanae (B; 650/7615) Hydrangeales (B; 42/485) Hydrangeaceae (C; 17/250) TropMont-temp As(-Sak) C sE H Me(-Bis) N(BrC, NY) S(-Pat, Chile) Philadelphoideae (C; 7/135) SubtrMont-temp Euras(-Jap)-wMal; Penn & BrC-nwMex Kirengeshomoideae (C; 1/1) (Kirengeshoma) Temp Chi, Kor, Jap

CLASSIFICATION AND GEOGRAPHY OF THE FLOWERING PLANTS

277

Hydrangeoideae (C; 9/114) TropMont-temp e, seAsia(-Sak)-Bis; H; New York & Wash-Andes(-Pat) Escalloniaceae (C; 15/200) TropMont--cool temp Af(-CapeP) eAs(-Jap) eAu(Tas) C(-Mex) G I Me(--Sol) eN P(Fern) S(Andes-Fuegia) Z Escallonioideae (incl. Corokia but excl. Ixerba, Griselinia & Carpodetus) (C; 10/165) TropMont--eool temp Af seAs eAu(-Tas) C(--CRica) G I(-Masc) Me(Sol) P(-Fern, Poly) S(Andes--Fuegia) Z Iteoideae (2/22) (Choristylis, Itea) TropMont-temp Af(-CapeP); se, eAs(Jap)-wMal; NJer & Ill-Ha & Tex Tribeloideae (1/1) (Tribeles) Temp SAmer(-Fuegia) Phyllonomoideae (1/8) (Phyllonoma) TropMont Mex-Peru Pterostemonoideae (1/2) (Pterostemon) TropMont Mex Griseliniaceae (B; 1/6) (Griselinia) Temp Z & sSAmer Alseuosmiaceae (C; 3/12) (Alseuosmia, Crispiloba, Wittsteinia [incl. Memecylanthus, Pachydiscus,Periomphale])TropMont-temp eAu-NGu; NCal-Z Montiniaceae (incl. Kaliphora, Melanophylla) (A; 4/13) Trop--warm temp Af(Cape); Ma Brexiaceae (3/11) (Brexia, Ixerba, Roussea) Trop--warm temp eAf-Ma, Seyc & Masc; nZ Columelliaceae (C; 1/4) (Columellia) Andean Col-Bol Desfontainiaceae (B; 1/1) (Desfontainia) TropMont--cool temp CRica-Fuegia Cornales (B; 92/1285) Vitineae (B; 13/735) Vitaceae (C; 13/735) Trop--temp Aft--Cape) As(-Sak) At(Ber) Au C E G I(Masc) Ma Me(-Fiji) N(--Que, Wash) P(Car, Gal) S(--Chile) Wind Vitoideae (12/700) Trop--temp Af(--Cape) As(--Sak) At(Ber) Au C E G I(Masc) Ma Me(-Fiji) N(-Que, Wash) P(Car, Gal) S(--Chile) Wind Leeoideae (1/34) (Leea) Trop Afse, eAs(-China) Au(neQld) G I(-Masc) Ma Me(-Fiji) P(Car) Gunnerineae (B; 1/35) Gunneraceae (C; 1/35) (Gunnera) TropMont-temp Af(--Cape) Au(Tas) C(Mex) H Ma Me(Phil-NHeb) O P(Fern) S(-Fuegia) Z Haloragineae (B; 9/100) Haloragaceae (C; 9/100) Subcosm Aft-Cape) seAs(-Kam) At(Az, Ber) Au C E(-Ice) G I(-Masc) Ma Me(Sol) N(--Green) O P(-Rapa, Fern) S(-Fuegia) Wind Z Cornineae (C; 11/115) Cornaceae (C; 6/78) TropMont-bor eAfr As(-Kam) C E(-Ice) Me N(--Green)

S(-Bol)
Davidioideae (1/1) (Davidia) Temp sw & cChi Nyssoideae (2/6) (Camptotheca, Nyssa) TropMont-temp Him & AssamChi, Java 8: Born; Maine & Ont-Pan Mastixioideae (2/25) (Diplopanax, Mastixia) TropMont seAs-Sol Cornoideae (1/46) (Cornus) TropMont eAf; temp--Arct Euras(Ice--Kam-Chi); IropMont-Arct Green & Alas--Bol Curtisiaceae (C; 1/l) (Curtisia) Subtr-temp SAf (Transv--Cape) Alangiaceae (C; 1/19) (Alangium) Trop Af-Ma & Comoro Is; se, eAs(-Mac)Fiji, eAu & NCal

278

THE BOTANICAL REVIEW

Garryaceae (C; 1/12) (Garrya) TropMont-temp Wash-Pan; Jam & Cuba Aucubaceae (C; 1/3) (Aucuba) Temp Him & Assam-Kor & Jap Aralidiaceae (C; 1/1) (Aralidium) Trop sThailand-Sum & Born Eucommiaceae (C; 1/1) (Eucommia) Temp Chi Icacinaceae (excl. Metteniusa) (A; 56/300) Trop--warm temp Af(-Cape) se, eAs(-Jap) eAu C(-Mex) G I(-Masc) Ma Me(-Fiji) P(-Tonga) S(-Chile) Wind Z Cardiopteridaceae (A; 1/2) (Peripterygium) Trop Assam, sChi-Bis; neQld Pittosporales (B; 14/195) Pittosporaceae (C; 10/150) Trop-temp Af(-Cape) e, seAs(-Jap) At(Mac) Au G H I(-Masc, Seyc) Ma Me(-Fiji) P(-Poly) Z Byblidaceae (B; 1/2) (Byblis) Trop--temp Au, NGu Tremandraceae (B; 3/43) (Platytheca, Tetratheca, Tremandra) Temp Au (esp. sw) Araliales (C; 462/4260) Helwingiaceae (C; 1/5) (Helwingia) SubtrMont-temp Him & Assam-Chi & Jap Torricelliaceae(C; 1/3) (Torricellia) SubtrMont-tempHim(wNepal)& nBurmasChi Araliaceae (C; 50/1150) Trop--bor Af(--Cape) As(-Sak) At(Mac) Au C E G H I(-Masc, Seyc) Ma Me(-Fiji) N(-Green) O P(-Marq) S(-Fuegia) Wind Z Hydrocotylaceae (C; 45/300) Subcosm Af(--Cape)As(-Jap) At(Can, Berm) Au C E(-Ice) G I(Masc, Seyc) Ma Me(-Fiji) N(-Nf, Wash) O P(-Marq) S(Fuegia) Wind Z Apiaceae (C; 365/2800) Subcosm Af(-Cape) As(-Kam) At(Mac) Au C E(-Ice) G H Ma Me(-NGu) N(--Green) O P(-Fern, Rapa) S(-Fuegia) Wind Z Saniculoideae Temp Aft-Cape) As(-Sak) At(-Az) Au C E H wMe N(-Que, BrC) P(Fern) S(-Chile) Wind Z Apioideae Subcosm Af(-Cape) As(-Kam) At(Mac) Au C E(-Ice) G H Ma Me(-NGu) N(--Green) O P(-Rapa, Fern) S(-Fuegia) Wind Dipsacales (C; 40/1390) Caprifoliaceae (C; 12/450) TropMont-bor Euras(Ice-nAf-Kam)-Sum & Phil; Green-Mex Adoxaceae (C; 5/243) TropMont-bor eAfr As(-Kam) At eAu C E(-nAfr) Me(Phil) N(-Alas) P(Bon) S(--Chile) Adoxoideae (incl. Sambucus) (C; 4/43) TropMont-bore Af As(-Kam) At(Mac) eAu C E(-nAfr) Me(-Phil) N(-Alas) P(Bon) S(--Chile) Viburnoideae (1/200) (Viburnum) TropMont-bor As(-Sak) At(Can, Mad) C E(-nAfr) Me(-Phil) N(-Alas) P(Bon) S(Andes--Arg)Wind Valerianaceae (C; 13/400) TropMont-bor, Antarct Af(--Cape)As(-Sak) At(Mac) C E(Ice--nAfr)N(-Alas) S(Andes--Fuegia)Wind Triplostegiaceae (C; 1/2) (Triplostegia) TropMont-warm temp se & eAs--Celebes & NGu Dipsacaceae (C; 8/280) TropMont-tempAf(--Cape)As(-Manc) At(Mac) E(Ice-nAfr) Morinaceae (C; 1/17) (Morina) Temp Balkans & Israel-Chi Asteranae (incl. Campanulanae) (B; 1251/21,500) Asterales (C; 1164/19,125) Calyceraceae (C; 4/40) Subtr-temp SAmer(Bol & sBraz-Fuegia)

CLASSIFICATION AND GEOGRAPHY OF THE FLOWERING PLANTS

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Asteraceae (C; 1160/19,085) Subcosm Aft--Cape) As(-Kam) At(Mac, SHel) Au C E(Ice-nAfr) G H I(-Masc) Ma Me(-Fiji) N(--Green) O P(-Marc0 S(-Fuegia) Wind Z Lactucoideae (Cichorioideae) (340/6370) Trop--temp Af As(-Kam) At(Mac) Au C E(Ice-nAfr) G I Ma Me(-Fiji) N(--Green) P(Fern) S(-Fuegia) Wind Z Barnadesieae (9/95) Trop SAmer Mutisieae (81/800) Trop-temp Af(--Cape) As(China) C sN(Cal-wTex) S(-Fuegia) Tarchonantheae (2/27) (Brachylaena, Tarchonanthus)Trop Af-SAf; Masc Cardueae (incl. Carlineae, Echinopsideae, Eremothamneae, Gundelieae) (80/2610) Trop--bor Af As(-Kam) At(Mac) C E(Ice--nAfr) I(Maur, Seyc) Me(Phil) N(--Green) P(Bon, Fern) S(--Chile) Vernonieae (incl. Stokesia, Trichospira) (70/1500) Trop--temp Af As(Jap) Au C I(Masc, Seyc) Ma Me(-Fiji) N(-Ont, Sask) P(Gal, Rev) S(-Chile) Wind Liabeae (15/160) Trop Mex-Peru & Bol; Wind Lactuceae (Cichorieae) (70/1000) Trop--bor Af As(-Kam) At(Mac) Au C E(Ice-Afr) I (Maur, Seyc) Me(-NGu) N(--Green) P(Bon, Fern) S(Fuegia) Wind Arctoteae (excl. Ursinia) (13/180) Temp Af(--Cape); sw, cAs Asteroideae (820/12,715) Subcosm Af(--Cape) As(-Kam) At(Mac) Au C E(Ice-nAfr) G H I(Masc) Ma Me(-Fiji) N(-Green) O P(-Marc0 S(Fuegia) Wind Z Astereae (135/2500) Subcosm Af(--Cape) As(-Kam) At(SHel) Au C E(Ice-nAfr) G H I(Masc, Seyc) Ma Me(-Fiji) N(--Green) O P(--Sam) S(Fuegia) Wind Z Anthemidae (incl. Ursinia, Cotula) (107/1650) Subcosm Af(-Cape) As(Kam) At(Mac) Au C E(-Ice) G H Ma Me(-NGu) N(--Green) O P(Bon) S(-Fuegia) Z Inuleae (incl. Gnaphalieae) (185/2050) Subcosm Af As(-Kam) At(Mac) Au C E(-Ice) G H I(-Masc) Ma Me(-Fiji) N(--Green) O P(-Rapa) S(Fuegia) Wind Z Senecioneae (incl. Blennosperma) (100/2000) Subcosm Af(--Cape) As(Kam) At(Mac) Au C E(-Ice) G I(Masc, Seyc) Ma Me N(-Green) O P(-Poly, Fern) S(-Fuegia) Wind Z Calenduleae (7/110) Temp Af(-Cape); Mac & Med-sw, cAs; Fuegia Eupatorieae (60/2000) Subcosm Af As(--Sak) At Au C E G I(-Masc, Seyc) Me(-Fiji) N(-Nf, BrC) O P(-Tonga) S(--Chile) Wind Heliantheae (incl. Arnica, Bahiinae, Gaillardiinae, Helenieae, Madieae) (177/1630) Subcosm Af As(-Sak) Au C H I(-Masc) Ma Me(-Fiji) N(-Green) P(--Gal) S(-Fuegia) Wind Coreopsideae (32/535) Trop--temp AfAs At(s) Au C E H Me N P(-Poly) S Wind Tageteae (17/240) Trop--warm temp Fla to Cal-sAmer; Gal Campanulales (C; 87/2370) Menyanthaceae (C; 5/40) Subcosm Af(-Cape) As(-Kam) Au(-Tas) C E(Ice-nAfr) G I(-Maur) Ma Me(-Fiji) N(--Green) S(-Arg) Wind Z

280

THE BOTANICAL REVIEW

Pentaphragmataceae (C; 1/30) (Pentaphragma) Trop Burma & Chi-NGu Sphenocleaceae (C; 1/2) (Sphenoclea) Trop wAfr (poss. introd, to Ma; sAsNGu & neQld) Campanulaceae (C; 65/2000) Subcosm Af(-Cape) As(-Kam) At(Mac) Au(Tas) C E(Ice--nAfr) G H I(-Masc) Ma Me(-Fiji) N(--Green) O P(-Marq) S(-Fuegia) Wind Z Campanuloideae (34/820) Subcosm Af(-Cape) As(-Kam) At(Mac) Au(-Tas) C E(Ice--nAfr) G I Ma Me(-NGu) N(-Green) P(-Tonga) S(-Chile) Wind Z Cyphioideae (1/50) (Cyphia) Trop-warm temp Cape Verde Islands-SAf(CapeP) Lobelioideae (incl. Cyphocarpus)(30/l 130) Subcosm Af(--Cape)w, se, eAs(Kam) At(Mac) Au C wE(-nAfr) H I(-Masc) Ma Me(-Fiji) N(-sAlas) O P(-Marq) S(-Fuegia) Wind Z Goodeniaceae (C; 15/300) Trop--temp Af(-CapeP) se, eAs(-Jap) At(Ber) Au(Tas) C(-Mex) G H I(-Maur) Ma Me(-Fiji) seN(Ha, Tex) P(-Marq, Gal) S(-Chile) Wind Z Goodenioideae (12/210) Trop-temp Af(-CapeP) se, eAs(-Jap) At(Ber) Au(Tas) C(-Mex) G H I(-Masc, Seyc) Ma Me(-Fiji) seN(Ha, Tex) P(-Marq, Gal) S(-Chile) Wind Z Dampieroideae (3/88) (Anthotium, Dampiera, Leschenaultia) Temp Au(Tas) Solananae (C; 296/7735) Solanales (C; 296/7735) Solanineae (C; 141/4755) Solanaceae (C; 76/2900) Subcosrn Af As(-Sak) At(Mac) Au C E(-nAfr) G H I(-Masc) Ma Me(-Fiji) N(-Que, Wash) P(-Marq) S(-Arg, Chile) Wind Z Solanoideae (55/2400) Subcosm Af As(-Sak) At(Mac) Au C E(-nAfr) G H I(-Masc) Ma Me(-Fiji) N(-Que, Ore) P(-Marq) S(-Pat) Wind Z Cestroideae (20/485) Trop-temp swAfAu C G H N(-Wash) P(-Marq, Rev) S(-Pat) Sclerophylacoideae (l/12) (Sclerophylax) Subtr Par-Urug & Arg Duckeodendraceae (B; 1/l) (Duckeodendron) Trop Amaz Goetzeaceae (B; 4/5) Trop Cuba-PRico Nolanaceae (C; 1/18) (Nolana) Trop-warm temp Peru-Chile; Gal Convolvulaceae (C; 59/1830) Subcosm Af As(-Sak) At(Mac) Au C E(-nAfr) G H I(-Masc) Ma Me(-Fiji) N(-Que, BrC) P(-Marq) S(-Arg) Wind Z Humbertioideae (1/1) (Humbertia) Trop Ma Dichondroideae (2/9) (Dichondra, Falkia) Temp--trop Af(-Cape) se, eAs(Jap) At(Ber) Au C G I(Masc) Ma Me(-Phil) N(Va-Cal) P(Gal, Fern) S(-Arg) Wind Z Convolvuloideae (55/1650) Subcosm Af As(-Jap) At(-Mac, Ber) Au C E(hAft) G H I(-Masc, Seyc) Ma Me(-Fiji) N(-Que, BrC) P(-Marq, Gal) S(-Arg) Wind Z Cuscutoideae (1/170) (Cuscuta) Subcosm Af(--Cape) As(-Sak) At(Mac) Au C E G H Me(-Fiji) N(-NS, Wash) P(-Poly, Gal) S(-Arg) Wind Z Boraginineae (C; 139/2660) Hydrophyllaceae (C; 18/250) Trop--bor Af(--CapeP)se, eAs At(Ber) Au(neQld) C H Ma Me(-Phil, Sul) N(--Green) P(Gal) S(-Fuegia) Wind

CLASSIFICATION AND GEOGRAPHY OF THE FLOWERING PLANTS

281

Boraginaceae (C; 117/2400) Subcosm Af(--CapeP) As(-Kam) At(Mac) Au C E(Ice--nAfr) H I(-Masc) Ma Me(-Fiji) N(-Green) O P(-Marq) S(-Fuegia) Wind Z Ehretioideae (11/145) Trop--subtr Af(-CapeP) se, eAs(-Jap) neAu C I(Masc, Seyc) Ma Me(-Sol) wN(-Wash) P(Gal) S(--Chile) Wind Cordioideae (4/255) Trop--warm temp Af(-CapeP) seAs neAu C G H I(Masc, Seyc) Ma Me(-Fiji) seN(sFla) P(-Marq) S(--Chile) Wind Heliotropoideae (incl. Ixorhea, Nogalia) (6/430) Subcosm AfAs(-Jap) A(Ber, Mac) Au C E G H I(-Masc, Seyc) Ma Me(-Fiji) sN(-Del, Ia) P(-Marq, Gal) S(--Chile) Wind Boraginoideae(95/1570) Subcosm Af(-Cape) As(-Kam) At(Mac) Au C E(IcenAfr) I(-Masc, Seyc) Ma Me(-NGu) N(--Green) O P(Guad) S(-Fuegia) Z WeUstedioideae (1/2) (Wellstedia) Subtrop Namibia; Socotra Hoplestigmataceae (B; 1/2) (Hoplestigma) Trop wAfrica (Gabon-Cameroon) Lennoaceae (C; 2/4) (Lennoa, Pholisma) Trop-warm temp Cal & Anz-Col & Ven Tetrachondraceae (A; 1/2) (Tetrachondra) Cool temp Z-Pat & Fuegia Polemoniineae (C; 16/320) Polemoniaceae (incl. Cobaea) (C; 16/320) TropMont-Arct nEuras(Ice-Kam); Green & Alas-Andes & Fuegia; Wind Loasanae (A; 13/280) Loasales (C; 13/280) Loasaceae (C; 13/280) Trop-warm temp s, neAf-Arab; BCol-Chile & Arg; Ha-Wind; Gal; Marq Mentzelioideae (2/71) (Eucnide, Mentzelia) Trop--temp BrC-Chile; FlaWind; Gal Loasoideae (7/200) Trop--warm temp sAf(Nam-CapeP); Somalia-Arab; Mex-Arg; Wind; Marq Gronovioideae (4/8) Trop--warm temp Tex-Ariz & Mex-nwSAmer; Gal; Wind Myrtanae (C; 463/8120) Myrtales (C; 463/8120) Lythrineae (B; 301/4470) Lythraceae (C; 25/460) Subcosm Af(--Cape)As(-Jap) At(Mac) Au C E(-nAfr) G I(-Masc) Ma Me(-Fiji) N(-Que, Wash) P(-Poly, Gal) S(-Chile) Wind Lythroideae (22/450) Subcosm Af(-Cape) As(-Jap) At(Mac) Au C E(-nAfr) G I(-Masc) Ma Me(-Fiji) N(-Que, Wash) P(-Poly) S(-Chile) Wind Duabangoideae (1/3) (Duabanga) Trop seAs(-sChi)-NGu Sonneratioideae (1/4) (Sonneratia) Trop eAf seAs(-sChi) nAu G I(-Seyc) Ma Me(-NHeb) P(-wcPoly) Punicoideae (1/2) (Punica) Warm temp Balkans-wHim; Socot Alzateaceae (C; 1/2) (Alzatea) Trop CRica-Peru Rhynchocalycaceae (C; l/l) (Rhynchocalyx) Trop Natal Penaeaceae (C; 5/25) Temp SAf(-CapeP) Oliniaceae (C; 1/10) (Olinia) Trop Af-Cape; (St. Helena I-nat?) Trapaceae (B; 1/1-3) (Trapa) Trop-temp swAf-nAfr; Euras(-Manc)-NGu Crypteroniaceae (C; 3/10) (Axinandra, Crypteronia, Dactylocladus)Trop Cey, Assam & sChi-Born & Phil

282

THE BOTANICAL REVIEW

Melastomataceae (C; 244/3360) Trop-warm temp Af(-CapeP) se, eAs(-sJap) nAu C(-Mex) G I(-Masc) Ma Me(-Fiji) eN(-NS, Ia) P(-Soc) S(-Arg) Wind Melastomatoideae (237/2950) Trop--warm temp Af(-CapeP) se, eAs(-sJap) nAu C(-Mex) G I(-Masc, Seyc) Ma Me(-Fiji) eN(-NS, Wisc) P(-Soc) S(-Arg) Wind Memecyloideae (incl. Astronia) (7/410) Trop Af(-CapeP) sAs nAu C(-Mex) I(-Masc, Seyc) Ma Me(-Fiji) P(--Soc) S(-seBraz) Wind Combretaceae (C; 20/600) Trop--subtr Af(-CapeP) As At(Ber) neAu C(-Mex) G I(-Masc) Ma Me(-Fiji) seN(sFla) P(-Marq, Gal) S(-Arg) Wind Strephonematoideae (1/6) (Strephomena) Trop wAf Combretoideae (19/595) Trop-subtr Af(-CapeP) As At(Ber) nAu C(-Mex) G I(-Masc, Seyc) Ma Me(-Fiji) se N(sFla) P(-Marq, Gal) S(-Arg) Wind Onagrineae (C; 18/650) Onagraceae (C; 18/650) Subcosm Af(-Cape) As(-Kam) At(Mac, Ber) Au C E(Ice--nAfr) G I(-Masc, Seyc) Ma Me(-Fiji) N(-Green) O P(-Marq, Gal) S(-Fuegia) Wind Z Myrtineae (C; 144/3000) Myrtaceae (C; 144/3000) Trop-cool temp Af(-Cape) se, eAs(-sJap) At(Ber) Au C(-Mex) sE(Med) G H I(-Masc) Ma Me(-Fiji) seN(sFla) O P(-Marq) S(--Chile) W(-Fuegia) Z Heteropyxidoideae (2/2) (Heteropyxis, Psiloxylon) Trop-subtrop SAf(TransvNatal) & Masc Myrtoideae (incl. I.eptospermoideae) (142/3000) Trop--warmtemp Af(-Cape) se, eAs(-sJap) At(Ber) Au C(-Mex) sE(Med) G H I(-Masc, Seyc) Ma Me(-Fiji) seN(sFla) O P(-Marq) S(-Fuegia) Wind Z Gentiananae (C; 2135/31,935) Gentianales (Rubiales) (C; 1009/14,145) Loganiaceae (B; 22/545) Trop--warm temp Af(-CapeP) As(-Jap) Au C(-Mex) G H I(-Masc) Ma Me(-Fiji) seN(-Va, Ind) O P(-Marq) S(-Arg) Wind Z Loganioideae (18/480) Trop--warm temp Af(-CapeP) eAs(-Jap) Au C(-Mex) G H I(-Masc, Seyc) Ma Me(-Fiji) seN(-Va, Ind) O P(-Poly) S(-Arg) Wind Z Potalioideae (3/65) (Anthocleista, Fagraea, Potalia) Trop Af(-Transv) seAs nAu G I(-Masc) Ma Me(-Fiji) P(-Marq) S(Col-Braz) Plocospermatoideae (1/1) (Plocosperma, incl. Lithophytum) Trop sMexGuat Rubiaceae (C; 500--600/9000) Subscosm Af(--Cape)As(-Kam) At(Mac) Au C E(Ice-nAfr) G H I(-Masc) Ma Me(-Fiji) N(--Green) O P(-Marq) S(-Fuegia) Wind Z Cinchonoideae (incl. Hillia) Trop--warm temp Af(--Cape) As(-Jap) Au C G H I(Masc) Ma Me(-Fiji) N(-Que) P(-Soc) S(--Chile) Wind Antirrheoideae (Guettardoideae) Trop Af seAs neAu C(-Mex) G I(-Masc, Seyc) Ma Me(-Fiji) seN(sFla) P(-Marq, Cocos) S(-Arg) Wind Ixoroideae Trop Af(-CapeP) se, eAs(-Jap) neAu C(-Mex) G H I(-Masc, Seyc) Ma Me(-Fiji) seN(sFla) P(-Marq, Gal) S Wind Rubioideae (incl. Theligoneae, Henriquesiaceae) Subcosm Af(-Cape) As(Kam) At(Mac) Au(-Tas) C E(Ice-nAfr) G H I(-Masc, Seyc) Ma Me(Fiji) N(--Green) O P(-Marq, Fern) S(-Fuegia) Wind Z Dialypetalanthaceae (C; 1/1) (Dialypetalanthus) Trop Braz

CLASSIFICATION AND GEOGRAPHY OF THE FLOWERING PLANTS

283

Apocynaceae (C; 355/3700) Trop--temp Aft--Cape) As(-Jap) At(Mac) Au C E(-nAfr) G H I(-Masc, Seyc) Ma Me(-Fiji) N(-Alas) P(-Marq) S(--Chile) Wind Z Plumerioideae (incl. Cerbereae) (101/.9)Trop Af(-CapeP) seAs n, eAu(-Tas) C(-Mex) E(-nAfr) G H I(-Masc, Seyc) Ma Me(-Fiji) N(-NJ, Kan) P(Marq, Gal) S(--Chile)Wind Apocynoideae (87/?) Trop--temp Af(--CapeP) As(-Jap) At(-Az) Au C E(nAfr) G I Ma Me(-Fiji) N(-Alas) P(Bon) S(--Chile) Wind Z Periplocoideae (C; 45/200) Trop-temp Af(-CapeP) As At(Can) Au E(Med) G I Ma Me(-Phil) Secamonoideae Trop Aft-Cape) seAs nAu C(-Mex) G I(-Masc) Ma Me(NHeb) N(-Pa, Ind) S Wind Asclepiadoideae Trop-temp Aft-Cape) As At(Can) Au C E G I(-Masc) Ma Me(-Fiji) N(-NB, BrC) P(-Tonga, Gal) S(-Chile) Wind Gentianaceae (C; 80/900) Subcosm Aft-Cape) As(-Kam) At(Mac) Au C E(IcenAfr) G H I(-Maur) Ma Me(-Fiji) N(-Green) O P(East) S(-Fuegia) Wind Z Saccifoliaceae (B; 1/1) (Saccifolium) Trop GuayH (nBraz-sVen) Scrophulariales (Bignoniales) (C; 1126/17,790) Scrophulariineae (Bignonineae) (C; 789/10,870) Oleaceae (C; 29/600) Trop-temp Af(--Cape)As(-Sak) At(Mac) Au C E(-nAfr) G H I(-Masc) Ma Me(-Fiji) N(-Nf, BrC) P(-Marq) S(-Chile) Wind Z Jasminoideae (incl. Nyctanthes) Trop-temp Af As(-Jap) At(Mac) Au C E(nAfr) G I(-Masc, Seyc) Ma Me(-Fiji) swN(Cal-Tex) P(-Marq) S(--Chile) Oleoideae (incl. Hesperelaea) Trop--temp Aft--Cape) As(-Sak) At(Mac) Au C E(-nAfr) G H I(Masc) Ma Me(-Fiji) N(-Nf, BrC) P(-Tonga, Rev) S Wind Z Buddlejaceae (C; 10/150) Trop--temp Af(--CapeP) As(-Jap) C I(-Masc) Ma Me(-NGu) swN(-Utah, Cal) P(Mar) S(-Chile) Wind Stilbaceae (C; 6/13) Temp SAf(CapeP) Retzioideae (1/1) (Retzia) Temp SAf(Cape of Good Hope) Stilboideae (5/12) Temp SAf(CapeP) Bignoniaceae (excl. Paulownieae) (C; 113/800) Trop--warm temp Af(-CapeP) As(--China) Au(-Tas) C(-Mex) G I(-Masc, Seyc) Ma Me(Sol) sN(-NJ, Cal) P(Car) S(-Chile) Wind nZ Pedaliaceae (incl. Trapella) (C; 12/70) Trop--warm temp Af(Socot-CapeP); Ma; se, eAs(-Jap)--NGu Martyniaceae (C; 2/12) (Martynia, Proboscidea) Trop--warm temp Tex & CalArg; Wind Myoporaceae (excl. Leucophylleae, Oftia)(C; 3/135)(Bontia, Eremophila, Myoporum) Trop--temp se, eAs(-Jap) Au G H I(-Masc) Me(Mol-NGu) P(-Poly) nS(Sur-Ven, nat?) Wind Z Scrophulariaceae (incl. Schlegelieae = Paulownieae, Capraria, Jerdonia, Leucophylleae, Ofiia) (C; 220/3000) Subcosm Aft-Cape) As(-Kam) At(Mac) Au(-Tas) C E(Ice-nAfr) G H I(-Masc, Seyc) Ma Me(-Fiji) N(-Green) O P(-Marq, Gal) S(-Fuegia) Wind Z Scrophularioideae (incl. Selagineae)Subcosm Aft-Cape) As(-Kam) At(Mac) Au C E(-Ice) I Ma Me(-Fiji) N(--Green) O P(-Marq, Gal) S(-Fuegia) Wind Z Rhinanthoideae Subcosm Aft-Cape) As(-Kam) At(-Mac, Falk) Au(-Tas) C

284

THE BOTANICAL REVIEW

E(Ice-nAfr) I(-Masc, Seyc) Ma Me(-Bis) N(-Green) O P(Fem) S(-Fuegia) Wind Z Orobanchoideae (13/150) Trop--bor Af(-Cape) As(-Kam) swAu C E(-nAfr) I(Socot) Ma Me(-NGu) N(-Alas) P(Bon) S(Andes-Chile) Gesneriaceae (C; 126/2850) Trop-warm temp Af(--CapeP)se, eAs(-Jap) neAu C sE G H I Ma Me(-NHeb) P S(-Chile) Wind Z Gesnerioideae (54/1280) Trop Wind & Mex-Cocos I & nArg, Bol Coronantheroideae (9/20) Trop-temp Solomon Is & NCaI-Z, Lord Howe I & NSW; Chile & wArg Cyrtandroideae (63/1550) Trop--warm temp Af(-CapeP) e, seAs(-Jap) Au(neQld) C(-Mex) sE G H I(Masc) Ma Me(-NHeb) P(-Sam) S(-Peru) Globulariaceae (C; 2/30) (Globularia,Poskea)Subtr-temp Somalia-Socot; MacswAs; Sweden-nAfr Plantaginaceae (C; 3/220) (Bougueria, Littorella, Plantago) Subcosm Af As(Sib) At(Mac) Au C E(Ice-nAfr) H Ma Me(ANGu) N(-Green) O P(-Rapa) S(Andes--Fuegia) Wind Z Lentibulariaceae (C; 4/170) Subcosm Af As(-Kam) Au C E(Ice-nAfr) G I(Maur) Ma Me(-NGu) N(--Green) P(-Micr, Gal) S(-Fuegia) Wind Z Acanthaceae (C; 256/2770) Trop-warm temp Af(-Cape) As(-Manc) At(Mac) nAu C E(-nAfr) G I(-Masc) Ma Me N(-Que) P(-Rapa, Gal) S(-Chile) Wind Nelsonioideae (5/15) Trop Af As nAu C(-Mex) Ma wMe sN(-SC) P(Rev) S Wind Thunbergioideae (4/205) Trop Aft--Cape) seAs Ma Mendoncioideae (2/60) (Gilletiella, Mendoncia) Trop Af; Ma; sMex-Bol Acanthoideae Trop--temp Af As(-Manc) At(Mac) neAu C E(-nAfr) G I(Masc, Seyc) Ma Me(-Fiji) N(-Que) P(-Rapa, GaD S(-Chile) Wind Ruellioideae Trop-temp Af(-CapeP) se, eAs(-Jap) nAu C Ma Me(-Fiji) N(NJ) P(Mar, GaD S(-Par) Wind Callitrichaceae (B; 1/40-60) (Callitriche) Subcosm Af(-CapeP) As(-Kam) At(Mac) sAu C E(Ice-nAfr) G Ma Me(-NGu) N(-Green) O P(Fern) S(Fuegia) Wind Z Hippuridaceae (B; 1/3) (Hippuris) Temp-Arct nEuras(Ice-nAfr-Kam); Green & Alas-sCal; cChile-Fuegia Lamiineae (C; 302/8055) Verbenaceae (only Verbenoideae) (B; 36/1035) Trop-warm temp Af(-Cape) As(-Manc) At(-Mac, Ber) Au C E(-nAfr) G I Ma Me(-NHeb) N(-NS, BrC) P(-Gal) S(-Chile) Wind Z Phrymataceae (B; l/l) (Phryma) Temp eAs(Sib-Him); Que & Man-Fla & Olda Symphoremataceae (B; 3/24) (Congea,Sphenodesme,Symphorema) Trop seAs-Sum; neQld Nesogenaceae (B; 3/11) (Asepalum, Cyclocheilon, Nesogenes) Trop Af (Somalia, Eth-Tanz); swArab; Ma-Masc Is; Tuamotu Is Avicenniaceae (C; 1/14) (Avicennia) Trop Af(-CapeP) seAs At(Ber) n, eAu C(-Mex) G I(-Masc, Seyc) Ma Me(-NHeb) seN(Fla-Tex) P(-Mar, Gal) S(-Braz) Wind nZ Lamiaceae (B; 258/6970) Subcosm Af(-Cape) As(-Kam) At(Mac) Au C E(IcenAfr) G H I(Masc) Ma Me(-Fiji) N(-Green) P(-Marq) S(-Fuegia) Wind Z

CLASSIFICATION AND GEOGRAPHYOF THE FLOWERINGPLANTS

285

Chloanthoideae (excl. Spartothamnella, but incl. Prostanthereae, Tectona) (B; 17/240) Au, Indomalesia Teucrioideae (incl. Viticoideae, Caryopteridoideae, Monochileae, Spartothamnella, Tetraclea, Teucrium, Trichostema) (39/15 20) Subcosm Af(CapeP) s, eAs(-Jap) ne Au C E(Med) G H I(-Masc) Ma Me(-Fiji) N P (-Marq) S(-Urug) Wind Z Ajugoideae (incl. Garrettia, Wenchengia) (6/56) Temp Af, Euras-w Me-Au Scutellarioideae (incl. Renschia, Tinnea, Holmskioldia) (4/320) Subcosm Af(-Malawi) As(-Amur) Au C E(-nAfr) I(Masc) Ma Me(-NGu) N(Alas, Lab), S(-Fuegia) Wind Pogostemonoideae (6/88) Trop-temp Euras-Indomalesia, Au, Ma, Masc Lamioideae (53/1060) Subcosm Af(-Cape) As(-Kam) At(Mac) Au C E(Ice-nAfr) H I Ma Me(-NHeb) N(--Green) P(-Soc) S(-Fuegia) Wind Z Nepetoideae (Ocimoideae) (133/3685) Subcosm Af As(-Manc) At(Mac) nAu C E(-nAfr) G H I(-Masc) Ma Me(-Fiji) N(-NC, Cal) P(-Marq) S(-Arg) Wind SUBCLASS:MONOCOTYLEDONEAE(LILIIDAE) Lilianae (B; 1184/29,550) Liliales (B; 148/2675) Melanthiineae (C; 25/155) Melanthiaceae (incl. Aletris, Chionographideae, Lophiola, Narthecieae, Petrosavieae, Tofieldieae, Xerophylleae) (C; 23/150) Trop--Arct Ice-Kam-Born; Green-Wind; Guy & Ven-Peru Campynemataceae (C; 2/4) (Campynema, Campynemanthe) Trop NCal; Temp Tas Liliineae (B; 45/865) Alstroemeriaceae (C; 4/160) Trop-temp Mex & WInd-Arg & Chile Colchicaceae (incl. Anguillarieae, Iphigenieae) (C; 17/170) Trop--temp Af(Cape) w, c, seAs At(Mac) Au(-Tas) E(Med) Ma Me(NGu) Z Liliaceae (B; 22/485) Trop--Arct Euras(-nAfr)-NGu & eAu; Green--Guat Tricyrtidoideae (incl. Scoliopeae, Uvularieae) (8/40) Trol>-bor eAs(-Jap)NGu & eAu; Que-Fla & La Lilioideae (incl. Calochortus, Gageeae, Medeoleae, Tulipeae) (14/445) Temp-Arct Euras(nAfr-Kam); Green--Guat Trilliaceae (C; 2/50) (Paris, Trillium) Temp--bor Euras(Ice-Kam); Que & BrCFla & cCal Iridineae (C; 77/1655) Iridaceae (C; 77/1655) Subcosm Af(-Cape) As(-Kam) At(Mac) Au(-Tas) C E(-nAfr) H I(-Masc) Ma Me(-NGu) N(--Green) P(Fern) S(-Fuegia) Wind Z Isophysidoideae (1/1) (lsophysis) Temp Tas Nivenioideae (incl. Geosiris) (6/85) Trop--temp Af(--Cape); Ma Iridoideae (incl. Sisyrinchieae, Tigrideae, Mariceae) (42/695) Subcosm Af(Cape) As(-Kam) At(Mac, Ber) Au(-Tas) C E H I(-Masc) Ma Me(-NGu) N(-Green) P(Fern) S(-Fuegia) Wind Z Ixioideae (28/875) Trop--temp Af(Cape-Med & Mac)-sw, cAs Burmanniales (C; 17/145) Burmanniaceae (C; 15/135) Trop-warm temp Afse, eAs(-Jap) nAu(-Tas) C(-

286

THE BOTANICAL REVIEW

Mex) Ma Me(--Sol) se, cN P(Car) S Wind Z Burmannioideae (9/100) Trop--subtr Af(-Moz) se, eAs(-Jap) n, eAu C(-Mex) Ma Me(Sol) seN(-Va, Tex) P(Car) S(-Arg) Wind Thismioideae (6/35) Trop--temp AfAs(Ceyl) Au(Tas) C(Pan) Me(Mal-NGu) N(III) S(Braz, Ecu) nZ Corsiaceae (C; 2/9) (Arachnitis, Corsia) Trop NGu-Sol; neQld; Bol-Pat & Chile Asparagales (B; 266/5560) Asparagineae (B; 125/2590) Asparagaceae (B; 26/440) Trop--bor Af As(-Kam) At(Mac) nAu C(-Mex) E I(Masc) Ma Me(-NGu) N(-Green) S(--Chile) Convallarioideae (incl. Aspidistreae, Ophiopogoneae, Polygonatae) (19/110) Trop--bor Euras(-Kamchatka)-Phil; Green-Mex Asparagoideae (1/312) (Asparagus) Trop--temp Af; Ma & Masc; Eur(-nAfr & Mac); s & eAs(--Sak); nAu-NGu Ruscoideae (3/8) (Danae, Ruscus, Semele) Warm temp Med-swAs(-Iran); Mac(-Azores) Herrerioideae (3/9) (Clara, Herreria, Herreriopsis) Trop Ma; Trop--temp Braz-Urug & Chile Luzuriagaceae (B; 4/10) (incl. Drymophila) Trop--temp At(Falk) eAu G Me(Java-Fiji) S(Peru-Fuegia) Z Asphodelaceae (incl. Aloeae) (B; 15/795) Trop--temp AfAs At(Mac) Au C(nMex) E(espMed) I(-Masc) Ma O Z Aphyllanthaceae (incl. Anthericaceae: Caesieae, Hodgsonioleae, Johnsonieae, Leucocrinum, Simethideae, Thysanoteae) (34/620) Trop--temp Af(--CapeP) As(-Jap) Au C(-Mex) E G Ma Me(-NGu) swN(Tex-Ariz) S(Peru-Chile) Z Phormiaceae (B; 8/35) Trop--temp seAf seAs(--Chi) Au(-Tas) G H I(-Masc) Ma Me(-Fiji) O P(-Marq) S(Ven-Bol) Z Phormioideae (incl. Dianelleae) (B; 7/32) Trop-temp seAf seAs Au(-Tas) GH I(-Masc) Ma Me(-Fiji) O P(-Marq) S(Ven-Bol) Z(-Norf) Doryanthoideae (1/3) (Doryanthes) Subtrop NSW, eQld Tecophilaeaceae (incl. Wallerieae) (B; 5/20) Warm temp Af(--Cape); Bol & Chile; Cal Lanariaceae (C; 1/1) (Lanaria) Temp SAf (CapeP) Hemerocallidaceae (C; 1/16) (Hemerocallis) Temp--bor Euras (-Sak) Dracaenaceae (B; 9/230) Trop--temp Af seAs At(Mac) Au(-Tas) C(Mex-Pan) G H I(Masc, Seyc, Socot) Ma Me(-Fiji) sN O P S(-Fuegia) Z Dracaenoideae (incl. Sansevierieae) (2/130) (Dracaena, Sansevieria) Trop-warm temp Af seAs At(Mac) neAu C(Mex-Pan) G H I(-Masc, Socot) Ma P(Car) Nolinoideae (3/50) (Beaucarnea, Dasylirion, Nolina) Subtrop--warm temp sUSA (SC-Fla, Cal)--Mex Astelioideae (4/50) Trop--subAntarct AfAu(-Tas) G H I(-Masc, Seyc) Me(Fiji) O P(-Marq) S(Braz-Fuegia) Z Hanguanaceae (B; 1/1 or 2) (Hanguana) Trop seAs-NGu; nAu; Palau I Agavaceae (incl. only Agaveae, Yucceae) (C; 8/300) Trop--temp sUSA-Braz; Wind Yuccoideae (2/42) (Samuela, Yucca) Trop--temp NCar-Jam; Iowa-Utah & Cal--C Amer

CLASSIFICATION AND GEOGRAPHY OF THE FLOWERING PLANTS

287

Agavoideae (6/260) Trop-temp Utah-Sur & Braz; NCar-Wlnd Hostaceae (C; 1/10) (Hosta) Temp eAs, esp. Jap Blandfordiaceae (B; 1/4) (Blandfordia) Temp Qld-Tas Dasypogonaceae (B; 9/72) Trop-temp Au(-Tas) G Me(NGu-NBrit) Dasypogonoideae (1/3) (Dasypogon) Temp swW Au Calectasioideae (1/2) (Calectasia) Temp Au Lomandroideae (incl. Lomandreae, Xeroteae) (5/65) Trop-temp Au; NCal; NGu-NBrit Kingioideae (2/2) (Baxteria, Kingia) Temp swW Au Xanthorrhoeaceae (1/28) (Xanthorrhoea) Temp Au (incl. Tas) Amaryllidineae (B; 141/2970) Ixiolirionaceae (C; 1/4) (Ixiolirion) Temp sw & cAs--Chi Hyacinthaceae (=SciUoideae)(incl. Bowieae, Chlorogaleae,Hesperocallis, Hyacintheae, Massonieae, Scilleae) (C; 40/900) Temp Af(--Cape); Ma; Mac & Med-sEur & As(-Manc); Penn & BrC-Mex; Peru--Chile Alliaceae(C; 30/720) Trop--Arct AfAs(-Kam) At(Mac) C E(Ice-nAfr) I(Masc) N S(-Pat) Wind Agapanthoideae (2/31) (Agapanthus, Tulbaghia) Trop Af-temp SAf(-Cape) Allioideae(incl. Brodiaeeae,Milleae)(19/665) Temp--borAs(-Kam) At(Mac) C E(Ice-nAfr) I(Masc) N(-Alas) S(-Pat) Wind GiUiesioideae (9/25) Trop--temp Peru-Chile Amaryllidaceae (C; 50/860) Trop--temp Af(--Cape) s, eAs(-Jap) At(Can) nAu C E(-nAfr) G I(-Masc, Seyc) Ma Me(-Fiji) N(-Va, Ky) P(--Car, Sam) S(Pat) Wind Hypoxidaceae (B; 10/150) Trop--temp Af(--Cape) s, eAs(-Jap) Au C(-Mex) G I(-Masc, Seyc)Ma Me(Sol) N(-Maine, Man) P(-Mar, Gal) S(-Arg)Wind Z Velloziaceae (B; 8/250) Trop eAf(--CapeP); Ma; Arab; Pan-Arg Vellozioideae (2/125) (Vellozia, Nanuza) Pan-sBraz Barbacenioideae(incl. Talbotia, Xerophyta) (6/125) Trop SAmer(-Arg);trop eAf(Somalia--CapeP); Ma; Arab Cyanastraceae (C; 1/6) (Cyanastrum) Trop Af Eriospermaceae (C; 1/80) (Eriospermum) Temp SAf(-Cape) Dioscoreales (B; 17/1050) Philesiaceae (B; 2/2) (Lapageria, Philesia) Temp SAmer(-Fuegia) Ripogonaceae (C; 1/70) (Ripogonum) Subtr-temp eAu-NGu; Z Smilacaceae(C; 3/310) (Heterosmilax, Pseudosmilax, Smilax) Trop--temp Af(Natal) As(-Jap) At(Mac) e, sAu C E(Med) G H I(Masc) Ma Me(-Fiji) N(Que, Ore) P(-Micro, cPoly) S(-Arg) Wind Dioscoreaceae (C; 5/625) Trop--temp Af As(-Jap) At(Mac) nAu C E(-nAfr) G I(Masc) Ma Me(-Fiji) N(-NE, Minn) P(-Marq) S(-Pat) Wind Stenomeridoideae (2/6) (Avetra, Stenomeris) Trop Ma; Mal--Sum, Phil Dioscoreoideae (3/620) (Dioscorea, Rajania, Tamus) Trop--temp Af As(Jap) At(Mac) nAu C E(-nAfr) G I(Masc) Ma Me(-Fiji) N(-NE, Minn) P(-Marq) S(-Pat) Wind Trichopodaceae (C; 1/1) (Trichopus) Trop Cey-Mal Stemonaceae (incl. Pentastemona) (C; 4/32) Trop--warm temp se, eAs(-Jap)Bis; neQld; Ga & Ala-Fla Taccaceae (C; 1/10) (Tacca) Trop Af e, seAs(-sChi) nAu I(--Seyc)Ma Me(Fiji) P(Micr-Tonga) nS(-Braz)

288

THE BOTANICAL REVIEW

Orchidales (C; 736/20,120) Orchidaceae (C; 736/20,120) Subcosm Af(-Cape) As(-Kam) At(-Az) Au C E(Ice--nAfr) G H I(-Masc) Ma Me(-Fiji) N(-Green) O P(-Marq) S(-Fuegia) Wind Z Apostasioideae (2/18) (Apostasia, Neuwiedia)Trop--warm temp sJap & ChiNGu & neQld Cypripedioideae (4/100-150) Trop-bor Euras(-Kam)-Sol; Nf & Alas-Braz & Bol Neottioideae Trop--temp AfAs(-Kam) At(Ber) Au C E(Ice-nAfr) G I(-Masc, Seyc) Ma Me(-Fiji) N(--Green) O P(-Soc) S(-Chile) Wind Z Orchidoideae Subcosm Af As(-Kam) At(Mac) Au C E(Ice-nAfr) G H I(Masc) Ma Me(-Fiji) N(--Green) O P(-Marq) S(-Chile) Wind Z Epidendroideae (incl. Vandoideae) Trop--Arct Af As(-Kam) Au C E(-Ice) G I(-Masc, Seyc) Ma Me(-Fiji) N(-Green) P(-Marq) S(-Arg) Wind Z Hydatellanae (A; 2/8) Hydatellales (C; 2/8) Hydatellaceae (C; 2/8) (Hydatella, Trithuria) Temp Au(-Tas); Z Triuridanae (B; 8/80) Triuridales (C; 8/80) Triuridaceae (incl. Lacandonia, Peltophyllum) (8/80) Trop--warm temp Af se, eAs(-sJap) Au(neQld) C(-sMex) G I(-Seyc) Ma Me(-Fiji) P(Car) S(-Braz) Alismatanae (C; 57/450) Alismatales (C; 34/235) Butomaceae (C; 1/1) (Butomus) Temp Euras(-nAfr) Alismataceae (C; 16/100) Subcosm Af As(-Sak) At(Mac) Au C E(-nAfr) Ma Me(-Fiji) N S(-Pat) Wind Limnocharitoideae (3/1 l) (Butomopsis, Hydrocleys, Limnocharis) Trop Af seAs nAu C(-Mex) Me(-NGu) S(-Arg) Wind Alismatoideae (13/90) Subcosm Af(-Namib) As(-Sak) At(Mac) Au C E(nAfr) Ma Me(-NGu) N(-Alas) S(-Pat, Chile) Wind Hydrocharitaceae (C; 17/130) Subcosm Af(-CapeP) As(-Jap) At(Ber) Au C E G H I(-Masc, Seyc) Ma Me(-Fiji) N(-Que, Sask) P(-Soc) S(-Chile, Arg) Wind Hydrocharitoideae (incl. Najas, Vallisnerioideae) 04/125) Subcosm Af(Natal) As(-Jap) Au C E G I(-Seyc, Masc) Ma Me(Sol) N(-Que, Sask) P(Car) S(-Arg, Chile) Wind Thalassioideae (2/3) (Enhalus, Thalassia)Trop-subtr eAfseAs At(Bet) nAu C(e) G I(-Seyc) Ma Me(Sol) seN(Fla-Tex) P(-Micr, wcPoly) nS Wind Halophiloideae (1/4) (Halophila) Trop--subtr eAf(-CapeP) se, eAs(-Jap) At(Ber) Au C E(Med) G H I(-Masc, Seyc) Ma Me(-Fiji) seN(Ha, Tex) P(-Soc) nS Wind Potamogetonales (Zosterales) (C; 23/215) Aponogetonineae (C; 1/47) Aponogetonaceae (C; 1/47) (Aponogeton)Trop-warm temp Af(-Cape)-Ma & Comoros; Taiwan & seChi-NGu; n, eAu-NCal Potamogetonineae (C; 19/162) Scheuchzeriaceae (C; 1/1) (Scheuchzeria) Temp--bor Euras (-Kam); Green & Alas-N Jet, Indiana & nCal

CLASSIFICATION AND GEOGRAPHY OF THE FLOWERING PLANTS

289

Juncaginaceae (C; 5/17) Subcosm Af(--Cape) As(-Kam) At(Gough) Au C(Mex) E(Ice-nAfr) Ma Me(-NGu) N(--Green) O S(-Fuegia) Wind Z Juncaginoideae (C; 4/16) Subcosm Af(-Cape) As(-Kam) At(Gough) Au C(Mex) E(Ice-nAfr) Ma Me(-NGu) N(-Green) O S(-Fuegia) Wind Z Lilaeoideae (C; 1/1) (Lilaea) Subtr-temp BrC-Mex; Par--Chile & Arg Potamogetonaceae (C; 3/95) (Groenlandia, Potamogeton, Ruppia) Subcosm Af(--Cape) As(-Kam) At(Mac) Au C E(Ice-nAfr) G H I(-Masc, Seyc) Ma Me(-Fiji) N(--Green) O P(Tonga, Gal) S(-Fuegia) Wind Z Potamogetonoideae (2/80-90) (Groenlandia, Potamogeton) Subcosm Af(Cape) As(-Kam) At(Mac) Au C E(Ice-nAfr) G H I Ma Me(-Fiji) N(Green) O P(-Tonga, Gal) S(-Fuegia) Wind Z Ruppioideae (1/3-7) (Ruppia) Trop--temp Af As Au C E Ma Me N(--Que, BrC) P(-Tonga, Gal) S Wind Z Posidoniaceae (C; 1/9) (Posidonia)Temp Mediterranean-swAs; Au(WA-TasNSW) Cymodoceaceae (C; 5/16) Trop--warm temp eAf(-Natal) se, eAs At(Can, Ber) Au eC(-Mex) E(Med) G I(-Masc) Ma Me(-Fiji) seN(NC-Fla, Tex) P(Tonga) nS(-Braz) Wind Zannichelliaceae (C; 4/12) Subcosm Af(-Cape) As(-Jap) At(Can) Au(-Tas) C E(Ice-nAfr) I(Masc) Ma N(-Alas) S(-Fuegia) Z Zosteraceae (C; 3/18) (Heterozostera, Phyllospadix, Zostera) Subtr-bor Af eAs(-Sak) At(Ber) e, s, wAu C(Mex) neE(Ice-Med) Ma N(-Green) P(Guad) S(Chile) Z Aranae (C; 108/2530) Arales (C; 104/2500) Araceae (excl. Acorus) (C; 104/2500) Subcosm Af(-Cape) As(-Kam) At(Mac) Au C E(-nAfr) G H I(-Masc) Ma Me(-Fiji) N(-Alas) P(-Marq) S(-Arg) Wind Gymnostachydoideae (1/2) (Gymnostachys) Trop eAu Pothoideae (incl. Monsteroideae) (18/1000) Trop Af seAs neAu C(-Mex) G I(-Masc) Ma Me(-Phil, Fiji) P(-Soc, Cocos) S(-Arg) Wind Calloideae (incl. Philodendroideae) (40/800) Trop--bor Af(--Cape)As(-Kam) Au(neQld) C(-Mex) E I(-Masc) Ma Me(-Sol) N(-Alas, Que) P(Cocos) S(-Arg) Wind Coeocasioideae (15/205) Trop Af se, eAs(-sJap) nAu C(Mex-Pan) G H I Ma Me(-Fiji) P(-Marq) S Wind Lasioideae (incl. Anthurieae, Callopsideae, Orontieae, Symplocarpeae) (12/ 80) Trop--bor Af eAs(-Jap) C(-Mex) Me(-Sol) N(--Que, Alas) P(-Marq) S(-Peru) Wind Aroideae (incl. Pistia) (18/415) Trop--cool temp Af As(-Sak) At(Mac) Au(nQld) C(-Mex) E(-nAfr) I(-Seyc, Masc) Me(-Fiji) N(--Que) S(-Arg) Lemnaceae (C; 4/28) Subcosm Af(-Cape) As(-Kam) At(-Az) Au C E(-nAfr) G H I(-Masc) Ma Me(-Fiji) N(-Alas) P(Gal) S(-Fuegia) Wind Z Lemnoideae (2/14) (Lemna, Spirodela) Subcosm Af(--Cape) As(-Kam) At(Mac) Au C E G H I(Masc) Ma Me(-Fiji) N(-Alas) P S(-Fuegia) Wind Z Wolflioideae (2/14) (Wolffia, Wolffiella)Trop--temp AfAs(-Jap) Au C E Ma wMe N(-NE, Minn) S(-Pat) Wind Z

290

T H E B O T A N I C A L REVIEW

Cyclanthanae (B; 11/240) Cyclanthales (C; 11/240) Cyclanthaceae (C; 11/240) Trop seMex & Ant-Bol Carludovicoideae (10/240) Trop seMex & Ant-Andes(--cBol) Cyclanthoideae (1/1) (Cyclanthus) Trop Guat & Ant-sePeru Pandananae (B; 3/700) Pandanales (C; 3/700) Pandanaceae (C; 3/700) Trop Af seAs nAu G H I(-Masc, Seyc) Ma Me(-Fiji) P(-Marc0 Z Pandanoideae (2/600) (Pandanus, Sararanga) Trop Af seAs nAu G H I(Masc, Seyc) Ma Me(-Fiji) P(-Marq) Freycinetioideae (1/100) (Freycinetia)Trop seAs neAu G H Me(-Fiji) P(Marq) Z Arecanae (C; 200/2780) Arecales (C; 200/2780) Arecaceae (C; 200/2780) Trop--warm temp Af(-Natal) se, eAs(-Jap) At(--Can, CVerde) n, eAu(-Vic) C E(Med) G H I(-Masc, Seyc) Ma Me(-Fiji) sN(NC, Cal) P(-Poly) S(-Chile) Wind Z Coryphoideae (incl. Phoenix, Borassoideae) (39/395) Trop-warm temp Af(Natal) s, eAs(Arab-sJap) At(Can, CVerde) n, eAu(-Vic) C E(Med) H I(-Masc, Seyc) Ma Me(-Fiji) sN(-NC, Cal) P(-Poly) S(-seBraz) Wind Calamoideae (=Lepidocaryoideae) (22/665) Trop Af(-Natal) seAs(-Taiw) neAu(-NSW) C(Nic--Pan) Ma Me(-Fiji) P(--Sam) S(-Braz, Peru) Nypoideae (1/1) (Nypa) Trop Ceylon & sChi--Sol & Car; neQld Ceroxyloideae (11/180) Trop Mex, Fla & Wlnd-Bol; Juan Fernandez Is; Comoro Is & Ma-Masc Arecoideae (incl. Caryotoideae, Cocosoideae) (124/1525) Trop--subtr Af(Natal) seAs(-Ry) neAu C(-Mex) G I(-Masc, Seyc) Ma Me(-Fiji) N(sFla) P(-Bon, Poly) S(-Par, Arg) Wind nZ Phytelephantoideae (3/15) (Ammandra, Polyandra, Phytelephas) Trop Pan-Bol Commelinanae (C; 1192/21,850) Bromeliales (C; 51/1520) Bromeliaceae (C; 51 / 1520) Trop--subtr wAf; Va & Tex-Pat & Chile; Wind; Gal Bromelioideae (30/425) Trop Mex & Wlnd-Arg, Chile, & Juan Fernandez Is Pitcairnioideae (incl. Navia) (13/420) Trop--subtr wAf (Guinea); Tex--Chile; Wind Tillandsioideae (8/675) Trop--warm temp Va-Fla & Wind; sAriz-Pat; Gal Philydrales (Pontederiineae) (C; 23/135) Philydraceae (C; 3/6) (Helmholtzia, Philydrella, Philydrum) Trop--temp se, eAs(-s Jap)-NGu; e, swAu Pontederiaceae (C; 6/30) Trop--temp Af(-Namib) seAs(-Jap) nAu C(-Mex) Ma Me(-NGu) N(--Que, Wash) S(-Arg) Wind Haemodoraceae (C; 14/100) Temp-trop Af(--Cape) seAs Au C(-Mex) Ma Me(NGu) N(-NS) S(-Braz) Wind Typhales (C; 2/30) Typhaceae (C; 2/30) Subcosm Af(-Cape) As(-Kam) At(Ber) Au C E(-Ice) G I(-Masc) Ma Me(-Fiji) N(--Green) S(-Pat) Wind Z

CLASSIFICATION AND GEOGRAPHY OF THE FLOWERING PLANTS

291

Sparganioideae (1/14) (Sparganium) Temp--Arct As(-Kam) eAu C(nwMex) E(Ice--nAfr) Me(NGu) N(--Green) Z Typhoideae (1/15) (Typha) Subcosm Af(--Cape) As(-Jap) At(Mac, Ber) Au C E(-nAfr) G I(-Masc, Seyc) Ma Me(-Fiji) N(-Alas) S(-Pat) Wind Z(Nor0 Zingiberales (C; 92/1975) Musineae (C; 2/42) Musaceae (C; 2/42) (Ensete, Musa) Trop Af seAs Au(neQld) G Ma Me(--Sol) P(Car) Strelitziineae (C; 3/7) Strelitziaceae (C; 3/7) (Phenakospermum, Ravenala, Strelitzia) Trop eAf(CapeP); Ma; Guy-Ecu Lowiineae (C; 1/6) Lowiaceae (C; 1/6) (Orchidantha) Trop sChi-Born Heliconiineae (C; 1/250) Heliconiaceae (C; 1/250) (Heliconia) Trop sMex & Wlnd-Arg; Sum-Fiji & Samoa; NCal Zingiberineae (C; 54/1150) Zingiberaceae (C; 50/1000) Trop Af se, eAs(-Jap) nAu C(-Mex) G H I(-Masc, Seyc) Ma Me(-Fiji) P(-Marq) S(-Par) Wind Costaceae (C; 4/150) Trop Afse, eAs Au(neQld) C(-Mex) Ma Me(--Sol)P(Car) S(-sBraz, Par) Wind Marantineae (C; 31/520) Cannaceae (C; 1/50) (Canna) Trop--subtr Mex-Arg; SCar-Wind Marantaceae (C; 30/450) Trop--subtr Af se, eAs C(-Mex) Ma Me(-NHeb) seN(-SC) P(Mar) S(-Arg) Wind Commelinales (C; 70/2140) Xyridineae (C; 20/350) Rapateaceae (C; 16/80) Trop wAf; Pan-nSAmer (esp GuayH) Saxofridericioideae (8/40) Trop Pan & GuayH-Amaz Braz Rapateoideae (8/40) Trop wAf; GuayH-Amaz(-Peru) Xyridaceae (C; 4/270) Trop--temp Af(-Cape) se, eAs Au C(-Mex) G I(-Masc) Ma Me(-NGu) eN(-Nf, Minn) P(Car) S(-Arg) Wind Xyridoideae (2/250) (Achlyphila, Xyris) Trop--temp Af(-Cape) seAs Au C(Mex) G I(-Maur) Ma Me(-NGu) eN(-Nf, Minn) P(Car) S(-Arg) Wind Abolbodoideae (2/20) (Abolboda, Orectanthe) Trop Guy & Ven-Braz Commelinineae (C; 41/615) Commelinaceae (C; 40/605) Trop--temp Af(--Cape) s, eAs(--Sak) At(Ber) Au C(-Mex) G H I(-Masc) Ma Me(-Fiji) N P(-Marq) S(-Arg) Wind Cartonematoideae (C; 2/7) (Cartonema, Triceratella)Trop Af; nAu Commelinoideae (C; 38/600) Trop--temp Af(--Cape) s, eAs(-Sak) At(Ber) Au C(-Mex) G H I(-Masc, Seyc) Ma Me(-Fiji) N(-Me, Minn) P(-Marq) S(-Arg) Wind Mayacaceae (C; 1/ 10) (Mayaca)Trop--warm temp Nam; Mex-Arg; Va & Miss-Wind Eriocaulineae (C; 9/1175) Eriocaulaceae (C; 9/1175) Trop-temp Af se, eAs(-Sib) n, eAu C(-Mex) E(Ire) G I(-Masc) Ma Me(-Bis) eN(-Nt) P(Car) S(-Arg) Wind

292

T H E B O T A N I C A L REVIEW

Eriocauloideae (2/410) (Eriocaulon, Mesanthemum) Trop-temp Af e, seAs(Sib) n, eAu C(-Mex) E(Ire) G I(-Masc) Ma Me(-Bis) eN(-Nf) P(Car) S(-Urug) Wind Syngonanthoideae (2/200) (Philodice, Syngonanthus) Trop-subtr Af C(-Mex) Ma seN(-NC) S(-sBraz) Wind Paepalanthoideae (5/565) Trop-subtr Af C(-Mex) seN(-Va) S(-Arg) Wind Juncales (Cyperales) (C; 155/5620) Thurniaceae (C; 1/3) (Thurnia) Trop GuayH, Amaz Juncaceae (C; 8/300) Subcosm Af(--Cape)As(-Kam) At(Mac) Au C E(-Ice) G H I(-Masc) Ma Me(-NGu) N(--Green) O P(--Soc, Fern) S(-Fuegia) Wind Cyperaceae (C; 146/5315) Subcosm Af(-Cape) As(-Kam) At(Mac, Bet) Au C E(-Ice) G H I(-Masc, Seyc) Ma Me(-Fiji) N(--Green) O P(-Marq) S(Fuegia) Wind Z Mapanioideae (l 4/200) Trop Af(-Cape) seAs Au(-Tas) C(-Mex) G I(-Maur, Seyc) Ma Me(-Fiji) P(--Sam) S(-Braz) Wind Cyperoideae (57/2750) Subcosm Af(-Cape) As(-Kam) At(Mac) Au C E(Ice) G H I(-Masc) Ma Me(-Fiji) N(--Green) O P(-Marq) S(-Fuegia) Wind Z Sclerioideae (70/365) Trop--temp AfAs(-Jap) Au C G H I(-Maur, Seyc) Ma Me(-Fiji) N(-NE, Minn) P(-Sam, Gal) S(-Arg) Wind Caricoideae (5/2000) Subcosm Af(-Cape) As(-Kam) At(Mac) Au C E(-Ice) G H I(-Masc) Ma Me(-Fiji) N(--Green) O P(Marq) S(-Fuegia) Wind Z Poales (Restionales) (C; 700/10,450) Flagellariineae (C; 49/450) Flagellariaceae (excl. Hanguana) (1/4) (Flagellaria) Trop Af(-Natal) seAs Au(neQld) G I(-Masc, Seyc) Ma Me(-Fiji) P(-Sam) Joinvilleaceae (1/2) (Joinvillea) Trop Mal & Sum-Palawan-Fiji & Samoa; Car; NCal; H Restionaceae (C; 41/405) Trop-temp Af(-Cape) seAs Au(-Tas) Ma Me(MalNGu) sS(cChile) Z Restionoideae (40/400) Temp-trop Af(-Cape) seAs Au Ma Me(Mal-NGu) sS(cChile) Z Anarthrioideae (1/7) (Anarthria) Temp swWAu Ecdeiocoleaceae (2/2) (Ecdeiocolea) Temp swWAu Centrolepidaceae (C; 4/30-35) Trop-cool temp seAs(-sChi) Au(-Tas) Me(Phil, NGu) O S(Fuegia) Z Poineae (C; 650/10,000) Poaceae (C; 650/10,000) Subcosm Af(-Cape) As(-Kam) At(-Mac) Au C E(Ice) G H I(-Masc) Ma Me(-Fiji) N(-Green) O P(-Marq) S(-Ant) Wind Z Bambusoideae (incl. Anomochloeae, Arundinarieae, Ehrharteae, Olyreae, Parianeae, Streptochaeteae) (79/1040) Trop--temp Af(-CapeP) As(-Sak) Au C E G I(Maur) Ma Me N(-Que, BrC) P S(-Chile) Wind Oryzoideae (Oryzae, Zizanieae) (12/70) Trop--temp Af As Au C E G I Ma Me N(-Que, BC) P S Wind Arundinoideae (incl. Aristideae, Arundineae, Centotheceae, Micrairieae, Stipeae, Thysanolaeneae) (55/645) Subcosm Af(-CapeP) As(-Manc) At(Mac) Au C E(nAfr) G I(-Masc, Seyc) Ma Me(-Fiji) N(-Alas) O P(Marq, Fern) S(-Fuegia) Wind Z

CLASSIFICATION AND GEOGRAPHY OF THE FLOWERING PLANTS

293

Poiideae (incl. Agrostideae, Aveneae, Bromeae, Lygeae, Nardeae, Triticeae) (153/3275) Subcosm Aft-Cape) As(-Kam) At(Mac) Au C E(-Ice) G H I(Masc) Ma Me(-Fiji) N(-Green) O P(-Marq, Fern) S(-Fuegia) Wind Z Chloridoideae (incl. Eragrostideae, Muhlenbergia, Neyraudia, Pappophorineae, Triraphis) (145/1360) Trop-cool temp Aft-Cape) As At(Tris) Au C E G I(-Masc, Seyc) Ma Me(-Sol) N(-Alas) P(-Cook) S(-Arg) Wind Panicoideae (incl. Andropogoneae, Eriachne) (207/3290) Subcosm Aft-Cape) As(-Manc) At(Mac) Au C E G H I(-Masc, Seyc) Ma Me(-Fiji) N(-NS, Que) P(-Marq) S(-Arg) Wind

TAXA INCERTAE SEDIS

Acorus L., Acoraceae (3) Temp--bor Euras(-Sib); Que & Alta-Fla & Tex. Removed
from Araceae and Arales by Grayum (1987a, 1990).

Aextoxicon Ruiz. & Pav., Aextoxicaceae (1) Temp Chile. Removed from Euphorbiales.

Barbeya Schweinf., Barbeyaceae (1) Subtr Eth-Arab. Removed from Urticales. Behnia Didrichsen (1) SAf. Removed from Philesiaceae and Luzuriagaceae. See
Conover (1991).

Carpodetus J. R. & G. Forst., Carpodetaceae (10) Trop NGu; Temp Z. Removed


from Escalloniaceae.

Haptanthus Goldberg & Nelson S. (1) Trop Hond. Relationships still unknown. See
Goldberg and Nelson (1989).

Heteranthia Nesse et Mart. (1) Trop Braz. Probably in or near Solanaceae. Metteniusa Karst, Metteniusaceae (6) nCol & wVen-Peru. Removed from Alangiaceae or Icacinaceae, possibly related to latter.

Petermannia F. Muell. Peterrnanniaceae (1) Subtrop eAu. Removed from Dioscoreales. See Conover (1991).

Physena Nor. ex Thou., Physenaceae (2) Trop Ma. Removed from Capparaceae or
Passifloraceae.

Pteleocarpa Oliv. (1) Trop WMe. Removed from Ehretioideae of Boraginaceae (Veldkamp 1988).

Setchellanthus T. S. Brandegee (1) Temp Mex. Removed from Capparaceae but


probably related to that family.

Trichostephanus Gilg (1) W trop Af. Removed from Flacourtiaceae by Lempke (1988).

V. Acknowledgments
I am most grateful to all those botanists who have so generously supplied separates, books, specimens, and other information useful in compiling this review. I am especially indebted to my fellow phylogenists, Arthur Cronquist, Armen Takhtajan, and Rolf and Gertrud Dahlgren for fruitful discussions, generous sharing of references and literature, and not least for their friendship and encouragement. I am also indebted to the various curators who have allowed me the use of their herbaria or who have sent me specimens on loan or on exchange for study.

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Index to Scientific Names


Abatieae, 268

Abolboda, 291,323

Agapanthus, 287 Agathelpis, 308

Acalyphoideae, 267 Agavaceae, 229, 286, 294 Acanthaceae, 231,232, 233, 245, 284, 295, Agaveae, 286 299, 310 Agavoideae, 287 Acanthogilia, 239, 302 Agdestidaceae, 262 Acanthoideae, 284 Agdestis, 262 Acer, 228, 273, 326 Agrostideae, 293 Aceraceae, 324 Aizoaceae, 233, 241,245, 262, 296 Aceroideae, 242, 273 Aizooideae, 262 Acharia, 268 Ajugeae, 294 Achariaceae, 268 Ajugoideae, 243, 285 Achatocarpaceae, 229, 262 Akania, 273 Achatocarpus, 262, 320 Akaniaceae, 273 Achlyphila, 291 Alangiaceae, 277 Acoraceae, 228, 230, 243, 293 Alangium, 238, 277 Acorus, 289, 293, 307 Aldina, 304 Actinidia, 263 Aletris, 285 Actinidiaceae, 229, 263, 302, 327 Alismataceae, 246, 288, 318 Actinidioideae, 263 Alismatales, 229, 288, 299 Adenaria, 306 Alismatanae, 243, 288 Adoxaceae, 231,278 Alismatidae, 217, 219 Adoxoideae, 278 Alismatoideae, 288 Aegialitis, 266 Allantospermum, 272 Aegiceras, 265 Alliaceae, 233, 287, 324 Aesculus, 272 Altioideae, 287 Aetoxylon, 268, 315 Allocasuarina, 305 Aextoxicaceae, 242, 293 Alnus, 275 Aextoxicon, 293 Aloeae, 286 Afrostyrax, 266 Alooideae, 240, 320 Agapanthoideae, 287 Alseuosmia, 277

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