Angiosperm Phylogeny Group (APG) system

Transcription

1 Angisperm Phylgeny Grup (APG) system The Angisperm Phylgeny Grup, r APG, refers t an infrmal internatinal grup f systematic btanists wh came tgether t try t establish a cnsensus view f the taxnmy f flwering plants (angisperms) that wuld reflect new knwledge abut their relatinships based upn phylgenetic studies. As f 2010, three incremental versins f a classificatin system have resulted frm this cllabratin (published in 1998, 2003 and 2009). An imprtant mtivatin fr the grup was what they viewed as deficiencies in prir angisperm classificatins, which were nt based n mnphyletic grups (i.e. grups cnsisting f all the descendants f a cmmn ancestr). APG publicatins are increasingly influential, with a number f majr herbaria changing the arrangement f their cllectins t match the latest APG system. Angisperm classificatin and the APG Until detailed genetic evidence became available, the classificatin f flwering plants (als knwn as angisperms, Angispermae, Anthphyta r Magnliphyta) was based n their mrphlgy (particularly that f the flwer) and their bichemistry (what kinds f chemical cmpund they cntained r prduced). Classificatin systems were typically prduced by an individual btanist r by a small grup. The result was a large number f such systems (see List f systems f plant taxnmy). Different systems and their updates tended t be favured in different cuntries; e.g. the Engler system in cntinental Eurpe; the Bentham & Hker system in Britain (particularly influential because it was used by Kew); the Takhtajan system in the frmer Sviet Unin and cuntries within its sphere f influence; and the Crnquist system in the United States. Histry f classificatin The btanical term "Angisperm", frm the Ancient Greek αγγείον, angeín (receptacle, vessel) and σπέρµα, (seed), was cined in the frm Angispermae by Paul Hermann in 1690, as the name f that ne f his primary divisins f the plant kingdm. This included flwering plants pssessing seeds enclsed in capsules, distinguished frm his Gymnspermae, r flwering plants with achenial r schiz-carpic fruits, the whle fruit r each f its pieces being here regarded as a seed and naked. The term and its antnym were maintained by Carlus Linnaeus with the same sense, but with restricted applicatin, in the names f the rders f his class Didynamia. Its use with any apprach t its mdern scpe nly became pssible after 1827, when Rbert Brwn established the existence f truly naked vules in the Cycadeae and Cniferae, and applied t them the name Gymnsperms. Frm that time nwards, s lng as these Gymnsperms were, as was usual, reckned as dictylednus flwering plants, the term Angisperm was used antithetically by btanical writers, with varying scpe, as a grup-name fr ther dictylednus plants. In 1851, Hfmeister discvered the changes ccurring in the embry-sac f flwering plants, and determined the crrect relatinships f these t the Cryptgamia. This fixed the psitin f Gymnsperms as a class distinct frm Dictyledns, and the term Angisperm then gradually came t be accepted as the suitable designatin fr the whle f the flwering plants ther than Gymnsperms, including the classes f Dictyledns and Mnctyledns. This is the sense in which the term is used tday. In mst taxnmies, the flwering plants are treated as a cherent grup. The mst ppular descriptive name has been Angispermae (Angisperms), with Anthphyta ("flwering plants") a secnd chice. These names are nt linked t any rank.

2 The Wettstein system and the Engler system use the name Angispermae, at the assigned rank f subdivisin. The Reveal system treated flwering plants as subdivisin Magnliphytina (Frhne & U. Jensen ex Reveal, Phytlgia 79: ), but later split it t Magnlipsida, Lilipsida and Rspsida. The Takhtajan system and Crnquist system treat this grup at the rank f divisin, leading t the name Magnliphyta (frm the family name Magnliaceae). The Dahlgren system and Thrne system (1992) treat this grup at the rank f class, leading t the name Magnlipsida. The APG system f 1998, and the later 2003 and 2009 revisins, treat the flwering plants as a clade called angisperms withut a frmal btanical name. Hwever, a frmal classificatin was published alngside the 2009 revisin in which the flwering plants frm the Subclass Magnliidae. The internal classificatin f this grup has undergne cnsiderable revisin. The Crnquist system, prpsed by Arthur Crnquist in 1968 and published in its full frm in 1981, is still widely used but is n lnger believed t accurately reflect phylgeny. A cnsensus abut hw the flwering plants shuld be arranged has recently begun t emerge thrugh the wrk f the Angisperm Phylgeny Grup (APG), which published an influential reclassificatin f the angisperms in Updates incrprating mre recent research were published as APG II in 2003 and as APG III in Traditinally, the flwering plants are divided int tw grups, which in the Crnquist system are called Magnlipsida (at the rank f class, frmed frm the family name Magnliacae) and Lilipsida (at the rank f class, frmed frm the family name Liliaceae). Other descriptive names allwed by Article 16 f the ICBN include Dictylednes r Dictyledneae, and Mnctylednes r Mnctyledneae, which have a lng histry f use. In English a member f either grup may be called a dictyledn (plural dictyledns) and mnctyledn (plural mnctyledns), r abbreviated, as dict (plural dicts) and mnct (plural mncts). These names derive frm the bservatin that the dicts mst ften have tw ctyledns, r embrynic leaves, within each seed. The mncts usually have nly ne, but the rule is nt abslute either way. Frm a diagnstic pint f view the number f ctyledns is neither a particularly handy nr reliable character. Recent studies, as by the APG, shw that the mncts frm a mnphyletic grup (clade) but that the dicts d nt (they are paraphyletic). Nevertheless, the majrity f dict species d frm a mnphyletic grup, called the eudicts r triclpates. Of the remaining dict species, mst belng t a third majr clade knwn as the Magnliidae, cntaining abut 9,000 species. The rest include a paraphyletic gruping f primitive species knwn cllectively as the basal angisperms, plus the families Ceratphyllaceae and Chlranthaceae. Flwering plant diversity The number f species f flwering plants is estimated t be in the range f 250,000 t 400,000. The number f families in APG (1998) was 462. In APG II (2003) it is nt settled; at maximum it is 457, but within this number there are 55 ptinal segregates, s that th The diversity f flwering plants is nt evenly distributed. Nearly all species belng t the eudict (75%), mnct (23%) and magnliid (2%) clades. The remaining 5 clades cntain a little ver 250 species in ttal, i.e., less than 0.1% f flwering plant diversity, divided amng 9 families. The mst diverse families f flwering plants, in their APG circumscriptins, in rder f number f species, are: 2

3 Asteraceae r Cmpsitae (daisy family): 23,600 species; Orchidaceae (rchid family): 22,075 species; Fabaceae r Leguminsae (pea family): 19,400; Rubiaceae (madder family): 13,150; Paceae r Gramineae (grass family): 10,035; Lamiaceae r Labiatae (mint family): 7,173; Euphrbiaceae (spurge family): 5,735; Melastmataceae (melastme family): 5,005; Myrtaceae (myrtle family): 4,620; Apcynaceae (dgbane family): 4,555. In the list abve (shwing nly the 10 largest families), the Orchidaceae and Paceae are mnct families; the thers are eudict families. The genetic evidence which became available frm the 1980s nwards, analysed by phylgenetic methds, cnfirmed r clarified sme relatinships in existing systems f classificatin, but radically changed thers. The rapid increase in knwledge led t many prpsed changes; stability was "rudely shattered". This psed prblems fr all users f classificatin systems (including encyclpaedists). In the late 1990s, an infrmal grup f researchers frm majr institutins wrldwide came tgether under the title f the 'Angisperm Phylgeny Grup' r APG. Their intentin was t prvide a widely accepted and mre stable pint f reference fr angisperm classificatin. Their first attempt at a new system was published in 1998 (the APG system). As f 2010, tw revisins have been published, in 2003 (APG II) and in 2009 (APG III), each superseding the previus system. Eight researchers have been listed as authrs t the three papers, and a further 33 as cntributrs. A classificatin presents a view at a particular pint in time, based n a particular state f research. Independent researchers, including members f the APG, cntinue t publish their wn views n areas f angisperm taxnmy. Classificatins change, hwever incnvenient this is t users. Hwever, the APG publicatins are increasingly regarded as an authritative pint f reference. Sme examples: A significant number f majr herbaria, including Kew, are changing the rder f their cllectins in accrdance with APG. The influential Wrld Checklist f Selected Plant Families (als frm Kew) is being updated t the APG III system. In the USA, a recent phtgraphic survey f the plants f the USA and Canada is rganized accrding t the APG II system. In the UK, the latest editin f the standard flra f the British Isles (by Stace) is based n the APG III system. The previus editin was based n the Crnquist system. Peter F. Stevens, ne f the authrs f all three f the APG papers, maintains a web site, APweb, hsted by the Missuri Btanical Garden, which has been regularly updated since 2001, and is a useful surce fr the latest research in angisperm phylgeny which fllws the APG apprach. Principles The principles f the APG's apprach t classificatin were set ut in the first paper f 1998, and have remained unchanged in subsequent revisins. Briefly, these are: The Linnean system f rders and families shuld be retained. "The family is central in flwering plant systematics." An rdinal classificatin f families is prpsed as a "reference 3

4 tl f brad utility". Orders are cnsidered t be f particular value in teaching and in studying family relatinships. Grups shuld be mnphyletic (i.e. cnsist f all descendants f a cmmn ancestr). The main reasn why existing systems are rejected is because they d nt have this prperty, they are nt phylgenetic. A brad apprach is taken t defining the limits f grups. Thus f rders, it is said that a limited number f larger rders will be mre useful. Families cntaining nly a single genus and rders cntaining nly a single family are avided where this is pssible withut vilating the ver-riding requirement fr mnphyly. Althugh rders and families are used, abve this level, named clades are freely emplyed in additin. (Sme have later been given frmal names in a paper assciated with the 2009 revisin f the APG system.) The authrs say that it is "nt pssible, nr is it desirable" t name all clades in a phylgenetic tree; hwever, systematists need t agree n names fr sme clades, particularly rders and families, t facilitate cmmunicatin and discussin. APG I system The initial 1998 paper by the APG made angisperms the first large grup f rganisms t be systematically re-classified primarily n the basis f genetic characteristics. The paper explains the authrs' view that there is a need fr a classificatin system fr angisperms at the level f families, rders and abve, but that existing classificatins are "utdated". The main reasn why existing systems are rejected is because they are nt phylgenetic, i.e. are nt based n strictly mnphyletic grups (i.e. grups which cnsist f all descendants f a cmmn ancestr). An rdinal classificatin f flwering plant families is prpsed as a "reference tl f brad utility". The brad apprach adpted t defining the limits f rders resulted in the recgnitin f 40 rders, cmpared t, fr example, 232 in Takhtajan's 1997 classificatin. Other features f the prpsed classificatin include: Frmal, scientific names are nt used abve the level f rder, named clades being used instead. Thus eudicts and mncts are nt given a frmal rank n the grunds that "it is nt yet clear at which level they shuld be recgnized". A substantial number f taxa whse classificatin had traditinally been uncertain are given places, althugh there still remain 25 families f "uncertain psitin". Alternative classificatins are prvided fr sme grups, in which a number f families can either be regarded as separate r can be merged int a single larger family. Fr example, the Fumariaceae can either be treated as a separate family r as part f Papaveraceae. A majr utcme f the classificatin is the disappearance f the traditinal divisin f the flwering plants int tw grups, mncts and dicts. The mncts are recgnized as a clade, but the dicts are nt, with a number f frmer dicts being placed in separate grups basal t bth mncts and the remaining dicts, the eudicts r true dicts. The APG system f plant classificatin is the first, nw bslete, versin f a mdern, mstly mlecular-based, system f plant taxnmy that was published in 1998 by the Angisperm Phylgeny Grup. It was superseded in 2003 by a revisin, the APG II system, and then in 2009 by a further revisin, the APG III system. 4

5 The riginal APG system is unusual in being based, nt n ttal evidence, but n the cladistic analysis f the DNA sequences f three genes, tw chlrplast genes and ne gene cding fr ribsmes. Althugh based n mlecular evidence nly, its cnstituent grups prve t be supprted by ther evidence as well, fr example pllen mrphlgy supprts the split between the eudicts and the rest f the frmer dictyledns. The system is rather cntrversial in its decisins at the family level, splitting a number f lng-established families and submerging a number f ther families. It als is unusual in nt using btanical names abve the level f rder, that is, an rder is the highest rank that will have a frmal btanical name in this system. Higher grups are defined nly as clades, with names such as mncts, eudicts, rsids, asterids. The main grups in the system (all unranked clades) are: angisperms : mncts cmmelinids eudicts cre eudicts rsids eursids I eursids II asterids euasterids I euasterids II The APG system recgnises 462 families and 40 rders: these are assigned as fllws. In the beginning f each listing sme families r rders that are nt placed in a further clade: clade angisperms family Ambrellaceae family Austrbaileyaceae family Canellaceae family Chlranthaceae family Hydnraceae family Illiciaceae family Nymphaeaceae (+ family Cabmbaceae) family Rafflesiaceae family Schisandraceae family Trimeniaceae family Winteraceae rder Ceratphyllales family Ceratphyllaceae rder Laurales family Atherspermataceae family Calycanthaceae family Gmrtegaceae family Hernandiaceae family Lauraceae family Mnimiaceae family Siparunaceae rder Magnliales family Annnaceae 5

6 family Degeneriaceae family Eupmatiaceae family Himantandraceae family Magnliaceae family Myristicaceae rder Piperales family Aristlchiaceae family Lactridaceae family Piperaceae family Saururaceae clade mncts family Crsiaceae family Japnliriaceae family Nartheciaceae family Petrsaviaceae family Triuridaceae rder Acrales family Acraceae rder Alismatales family Alismataceae family Apngetnaceae family Araceae family Butmaceae family Cymdceaceae family Hydrcharitaceae family Juncaginaceae family Limncharitaceae family Psidniaceae family Ptamgetnaceae family Ruppiaceae family Scheuchzeriaceae family Tfieldiaceae family Zsteraceae rder Asparagales family Agapanthaceae family Agavaceae family Alliaceae family Amaryllidaceae family Anemarrhenaceae family Anthericaceae family Aphyllanthaceae family Asparagaceae family Asphdelaceae family Asteliaceae family Behniaceae family Blandfrdiaceae family Bryaceae family Cnvallariaceae family Dryanthaceae family Hemercallidaceae family Herreriaceae 6

7 family Hyacinthaceae family Hypxidaceae family Iridaceae family Ixiliriaceae family Lanariaceae family Laxmanniaceae family Orchidaceae family Tecphilaeaceae family Themidaceae family Xanthrrheaceae family Xernemataceae rder Discreales family Burmanniaceae family Discreaceae family Taccaceae family Thismiaceae family Trichpdaceae rder Liliales family Alstremeriaceae family Campynemataceae family Clchicaceae family Liliaceae family Luzuriagaceae family Melanthiaceae family Philesiaceae family Ripgnaceae family Smilacaceae rder Pandanales family Cyclanthaceae family Pandanaceae family Stemnaceae family Vellziaceae clade cmmelinids family Ablbdaceae family Brmeliaceae family Dasypgnaceae family Hanguanaceae family Mayacaceae family Rapateaceae rder Arecales family Arecaceae rder Cmmelinales family Cmmelinaceae family Haemdraceae family Philydraceae family Pntederiaceae rder Pales family Anarthriaceae family Centrlepidaceae family Cyperaceae family Ecdeicleaceae 7

8 family Ericaulaceae family Flagellariaceae family Hydatellaceae family Jinvilleaceae family Juncaceae family Paceae family Priniaceae family Restinaceae family Sparganiaceae family Thurniaceae family Typhaceae family Xyridaceae rder Zingiberales family Cannaceae family Cstaceae family Helicniaceae family Lwiaceae family Marantaceae family Musaceae family Strelitziaceae family Zingiberaceae clade eudicts family Buxaceae family Didymelaceae family Sabiaceae family Trchdendraceae (+ family Tetracentraceae) rder Prteales family Nelumbnaceae family Platanaceae family Prteaceae rder Ranunculales family Berberidaceae family Circaeasteraceae (+ family Kingdniaceae) family Eupteleaceae family Lardizabalaceae family Menispermaceae family Papaveraceae (+ family Fumariaceae and family Pteridphyllaceae) family Ranunculaceae clade cre eudicts family Aextxicaceae family Berberidpsidaceae family Dilleniaceae family Gunneraceae family Myrthamnaceae family Vitaceae rder Caryphyllales family Achatcarpaceae family Aizaceae family Amaranthaceae family Ancistrcladaceae family Asterpeiaceae 8

9 family Basellaceae family Cactaceae family Caryphyllaceae family Didiereaceae family Dincphyllaceae family Drseraceae family Drsphyllaceae family Frankeniaceae family Mlluginaceae family Nepenthaceae family Nyctaginaceae family Physenaceae family Phytlaccaceae family Plumbaginaceae family Plygnaceae family Prtulacaceae family Rhabddendraceae family Sarcbataceae family Simmndsiaceae family Stegnspermataceae family Tamaricaceae rder Santalales family Olacaceae family Opiliaceae family Lranthaceae family Misdendraceae family Santalaceae rder Saxifragales family Altingiaceae family Cercidiphyllaceae family Crassulaceae family Daphniphyllaceae family Grssulariaceae family Halragaceae family Hamamelidaceae family Iteaceae family Paeniaceae family Penthraceae family Pterstemnaceae family Saxifragaceae family Tetracarpaeaceae clade rsids family Aphliaceae family Crsssmataceae family Ixerbaceae family Krameriaceae family Picramniaceae family Pdstemaceae family Stachyuraceae family Staphyleaceae family Tristichaceae 9

10 family Zygphyllaceae rder Geraniales family Francaceae family Geraniaceae (+ family Hypsecharitaceae) family Greyiaceae family Ledcarpaceae family Melianthaceae family Vivianiaceae clade eursids I family Celastraceae family Huaceae family Parnassiaceae (+ family Lepurpetalaceae) family Stackhusiaceae rder Cucurbitales family Anisphylleaceae family Begniaceae family Criariaceae family Cryncarpaceae family Cucurbitaceae family Datiscaceae family Tetramelaceae rder Fabales family Fabaceae family Plygalaceae family Quillajaceae family Surianaceae rder Fagales family Betulaceae family Casuarinaceae family Fagaceae family Juglandaceae family Myricaceae family Nthfagaceae family Rhipteleaceae family Ticdendraceae rder Malpighiales family Achariaceae family Balanpaceae family Carycaraceae family Chrysbalanaceae family Clusiaceae family Dichapetalaceae family Erythrxylaceae family Euphrbiaceae family Euphrniaceae family Flacurtiaceae family Gupiaceae family Hugniaceae family Humiriaceae family Hypericaceae family Irvingiaceae 10

11 family Ixnanthaceae family Lacistemaceae family Linaceae family Malesherbiaceae family Malpighiaceae family Medusagynaceae family Ochnaceae family Pandaceae family Passiflraceae family Putranjivaceae family Quiinaceae family Rhizphraceae family Salicaceae family Scyphstegiaceae family Trigniaceae family Turneraceae family Vilaceae rder Oxalidales family Cephaltaceae family Cnnaraceae family Cunniaceae family Elaecarpaceae family Oxalidaceae family Tremandraceae rder Rsales family Barbeyaceae family Cannabaceae family Cecrpiaceae family Celtidaceae family Dirachmaceae family Elaeagnaceae family Mraceae family Rhamnaceae family Rsaceae family Ulmaceae family Urticaceae clade eursids II family Tapisciaceae rder Brassicales family Akaniaceae (+ family Bretschneideriaceae) family Bataceae family Brassicaceae family Caricaceae family Emblingiaceae family Gyrstemnaceae family Keberliniaceae family Limnanthaceae family Mringaceae family Pentadiplandraceae family Resedaceae family Salvadraceae 11

12 family Setchellanthaceae family Tvariaceae family Trpaelaceae rder Malvales family Bixaceae (+ family Diegdendraceae) family Cistaceae family Cchlspermaceae family Diptercarpaceae family Malvaceae family Muntingiaceae family Neuradaceae family Sarclaenaceae family Sphaersepalaceae family Thymelaeaceae rder Myrtales family Alzateaceae family Cmbretaceae family Crypterniaceae family Heterpyxidaceae family Lythraceae family Melastmataceae family Memecylaceae family Myrtaceae family Oliniaceae family Onagraceae family Penaeaceae family Psilxylaceae family Rhynchcalycaceae family Vchysiaceae rder Sapindales family Anacardiaceae family Biebersteiniaceae family Burseraceae family Kirkiaceae family Meliaceae family Nitrariaceae (+ family Peganaceae) family Rutaceae family Sapindaceae family Simarubaceae clade asterids rder Crnales family Crnaceae (+ family Nyssaceae) family Grubbiaceae family Hydrangeaceae family Hydrstachyaceae family Lasaceae rder Ericales family Actinidiaceae family Balsaminaceae family Clethraceae family Cyrillaceae 12

13 family Diapensiaceae family Ebenaceae family Ericaceae family Fuquieriaceae family Halesiaceae family Lecythidaceae family Marcgraviaceae family Myrsinaceae family Pellicieraceae family Plemniaceae family Primulaceae family Rridulaceae family Saptaceae family Sarraceniaceae family Styracaceae family Symplcaceae family Ternstremiaceae family Tetrameristaceae family Theaceae family Thephrastaceae clade euasterids I family Braginaceae family Plcspermataceae family Vahliaceae rder Garryales family Aucubaceae family Eucmmiaceae family Garryaceae family Oncthecaceae rder Gentianales family Apcynaceae family Gelsemiaceae family Gentianaceae family Lganiaceae family Rubiaceae rder Lamiales family Acanthaceae family Avicenniaceae family Bignniaceae family Buddlejaceae family Byblidaceae family Cyclcheilaceae family Gesneriaceae family Lamiaceae family Lentibulariaceae family Mypraceae family Oleaceae family Paulwniaceae family Pedaliaceae (+ family Martyniaceae) family Phrymaceae family Plantaginaceae 13

14 family Schlegeliaceae family Scrphulariaceae family Stilbaceae family Tetrachndraceae family Verbenaceae rder Slanales family Cnvlvulaceae family Hydrleaceae family Mntiniaceae family Slanaceae family Spencleaceae clade euasterids II family Adxaceae family Bruniaceae family Carlemanniaceae family Clumelliaceae (+ family Desfntainiaceae) family Eremsynaceae family Escallniaceae family Icacinaceae family Plysmaceae family Sphenstemnaceae family Tribelaceae rder Apiales family Apiaceae family Araliaceae family Aralidiaceae family Griseliniaceae family Melanphyllaceae family Pittspraceae family Trricelliaceae rder Aquifliales family Aquifliaceae family Helwingiaceae family Phyllnmaceae rder Asterales family Alseusmiaceae family Argyrphyllaceae family Asteraceae family Calyceraceae family Campanulaceae (+ family Lbeliaceae) family Carpdetaceae family Dnatiaceae family Gdeniaceae family Menyanthaceae family Pentaphragmataceae family Phellinaceae family Russeaceae family Stylidiaceae rder Dipsacales family Caprifliaceae family Diervillaceae 14

15 family Dipsacaceae family Linnaeaceae family Mrinaceae family Valerianaceae "+..." = ptinal seggregrate family, that may be split ff frm the preceding family. Families f uncertain psitin family Balanphraceae family Bnnetiaceae family Cardipteridaceae family Ctenlphnaceae family Cynmriaceae family Cytinaceae family Dipentdntaceae family Elatinaceae family Geisslmataceae family Hplestigmataceae family Kaliphraceae family Lepidbtryaceae family Lisscarpaceae family Lphpyxidaceae family Medusandraceae family Mettenusiaceae family Mitrastemnaceae family Paracryphiaceae family Pentaphylacaceae family Peridiscaceae family Plagipteraceae family Pttingeriaceae family Sladeniaceae family Strasburgeriaceae family Tepuianthaceae APG II system The secnd paper published by the APG presents an update t the riginal classificatin f The authrs say that changes have been prpsed nly when there is "substantial new evidence" which supprts them. The prpsed classificatin cntinues the traditin f seeking brad circumscriptins f taxa, fr example trying t place small families cntaining nly ne genus in a larger grup. The authrs say that they have generally accepted the views f specialists, althugh nting that specialists "nearly always favur splitting f grups" regarded as t varied in their mrphlgy. APG II cntinues and indeed extends the use f alternative bracketed" taxa allwing the chice f either a large family r a number f smaller nes. 15

16 Fr example, the large Asparagaceae family includes 7 bracketed families which can either be cnsidered as part f the Asparagaceae r as separate families. Sme f the main changes in APG II are: New rders are prpsed, particularly t accmmdate the 'basal clades' left as families in the first system. Many f the previusly unplaced families are nw lcated within the system. Several majr families are re-structured. In 2007, a paper was published giving a linear rdering f the families in APG II, suitable fr rdering herbarium specimens, fr exampl. The APG II system (2003) f plant classificatin is the secnd, nw bslete, versin f a mdern, mstly mlecular-based, system f plant taxnmy that was published in April f 2003 by the Angisperm Phylgeny Grup. It was a revisin f the first APG system, published in 1998, and was superseded in 2009 by a further revisin, the APG III system. APG II was published as: Angisperm Phylgeny Grup (2003). "An update f the Angisperm Phylgeny Grup classificatin fr the rders and families f flwering plants: APG II". Btanical Jurnal f the Linnean Sciety 141(4): Each f the APG systems represents the brad cnsensus f a number f systematic btanists, united in the APG, wrking at several institutins wrldwide. The APG II system recgnized 45 rders, five mre than the APG system. The new rders were Austrbaileyales, Canellales, Gunnerales, Celastrales, and Crsssmatales, all f which were families unplaced as t rder, althugh cntained in suprardinal clades, in the APG system. APG II recgnized 457 families, five fewer than the APG system. Thirty-nine f the APG II families were nt placed in any rder, but 36 f the 39 were placed in a supra-rdinal clade within the angisperms. Fifty-five f the families came t be knwn as "bracketed families". They were ptinal segregates f families that culd be circumscribed in a larger sense. The APG II system was influential and was adpted in whle r in part (smetimes with mdificatins) in a number f references. It was superseded 6½ years later by the APG III system, published in Octber f Main grups in the system (all unranked clades between the ranks f class and rder): angisperms : magnliids mncts cmmelinids eudicts cre eudicts rsids eursids I eursids II asterids euasterids I euasterids II Shwn belw is the classificatin in full detail, except fr the fifteen genera and three families that were unplaced in APG II. The unplaced taxa were listed at the end f the appendix in a sectin entitled "Taxa f Uncertain Psitin". 16

17 Under sme f the clades are listed the families that were placed incertae sedis in that clade. Thirtysix families were s placed. This means that their relatinship t ther members f the clade is nt knwn. paraphyletic grade basal angisperms family Ambrellaceae family Chlranthaceae family Nymphaeaceae (+ family Cabmbaceae) rder Austrbaileyales rder Ceratphyllales clade magnliids rder Canellales rder Laurales rder Magnliales rder Piperales clade mncts family Petrsaviaceae rder Acrales rder Alismatales rder Asparagales rder Discreales rder Liliales rder Pandanales clade cmmelinids family Dasypgnaceae rder Arecales rder Cmmelinales rder Pales rder Zingiberales clade eudicts family Buxaceae (+ family Didymelaceae) family Sabiaceae family Trchdendraceae (+ family Tetracentraceae) rder Prteales rder Ranunculales clade cre eudicts family Aextxicaceae family Berberidpsidaceae family Dilleniaceae rder Gunnerales rder Caryphyllales rder Santalales rder Saxifragales clade rsids family Aphliaceae family Geisslmataceae family Ixerbaceae family Picramniaceae family Strasburgeriaceae family Vitaceae rder Crsssmatales rder Geraniales 17

18 rder Myrtales clade eursids I family Zygphyllaceae (+ family Krameriaceae) family Huaceae rder Celastrales rder Cucurbitales rder Fabales rder Fagales rder Malpighiales rder Oxalidales rder Rsales clade eursids II family Tapisciaceae rder Brassicales rder Malvales rder Sapindales clade asterids rder Crnales rder Ericales clade euasterids I family Braginaceae family Icacinaceae family Oncthecaceae family Vahliaceae rder Garryales rder Gentianales rder Lamiales rder Slanales clade euasterids II family Bruniaceae family Clumelliaceae (+ family Desfntainiaceae) family Eremsynaceae family Escallniaceae family Paracryphiaceae family Plysmaceae family Sphenstemnaceae family Tribelaceae rder Apiales rder Aquifliales rder Asterales rder Dipsacales "+..." = ptinally separate family, that may be split ff frm the preceding family. In figure 1 are reprted the Cladgram f Angisperms. 18

19 Figure 1 - A cladgram shwing the relatinships, but excluding taxa nt placed within an rder. 19

20 APG III system The third paper frm the APG updates the system described in the 2003 paper. The brad utline f the system remains unchanged, but the number f previusly unplaced families and genera is significantly reduced. This requires the recgnitin f bth new rders and new families cmpared t the previus classificatin. The number f rders ges up frm 45 t 59; nly 10 families are nt placed in an rder and nly tw f these (Apdanthaceae and Cynmriaceae) are left entirely utside the classificatin. The authrs say that they have tried t leave lng-recgnized families unchanged, while merging families with few genera. They "hpe the classificatin will nt need much further change." A majr change is that the paper discntinues the use f 'bracketed' families in favur f larger, mre inclusive families. As a result, the APG III system cntains nly 415 families, rather than the 457 f APG II. Fr example, the agave family (Agavaceae) and the hyacinth family (Hyacinthaceae) are n lnger regarded as distinct frm the brader asparagus family (Asparagaceae). The authrs say that alternative circumscriptins, as in APG I and II, are likely t cause cnfusin and that majr herbaria which are re-arranging their cllectins in accrdance with the APG apprach have all agreed t use the mre inclusive families. In the same vlume f the jurnal, tw related papers were published. One gives a linear rdering f the families in APG III; as with the linear rdering published fr APG II, this is intended fr rdering herbarium specimens, fr example. The ther paper gives, fr the first time, a classificatin f the families in APG III which uses frmal taxnmic ranks; previusly nly infrmal clade names were used abve the rdinal level. The APG III system (2009) f plant classificatin is the third versin f a mdern, mstly mlecular-based, system f plant taxnmy that was published in 2009 by the Angisperm Phylgeny Grup, 6½ years after its predecessr, the APG II system was published. In Octber 2009, members f the Linnean Sciety prpsed an accmpanying frmal phylgenetic classificatin f all land plants, cmpatible with the APG III classificatin.[4] This was needed because btanists and phyclgists ften disagree n the taxnmic rank t which grups are assigned. The APG III system recgnized all f the 45 rders f the previus system, as well as 14 new nes. The rder Ceratphyllales was errneusly marked as a new rder, but it had been recgnized in bth f the previus APG systems. The newly recgnized rders were: Ambrellales, Nymphaeales, Chlranthales, Petrsaviales, Trchdendrales, Buxales, Vitales, Zygphyllales, Picramniales, Huerteales, Berberidpsidales, Escallniales, Bruniales, and Paracryphiales. The designatin f alternative "bracketed families" was abandned in APG III, because its inclusin in the previus system had been unppular. APG III recgnized 415 families, 42 fewer than in the previus system. Frty-fur f the 55 "bracketed families" were discntinued, and 18 ther families were discntinued as well. The discntinued bracketed families were: Illiciaceae, Alliaceae, Agapanthaceae, Agavaceae, Aphyllanthaceae, Hespercallidaceae, Hyacinthaceae, Laxmanniaceae, Ruscaceae, Themidaceae, Asphdelaceae, Hemercallidaceae, Kingdniaceae, Fumariaceae, Pteridphyllaceae, Didymelaceae, Tetracentraceae, Pterstemnaceae, Hypsecharitaceae, Francaceae, Memecylaceae, Lepurpetalaceae, Rhipteleaceae, Medusagynaceae, Quiinaceae, Malesherbiaceae, Turneraceae, Bretschneideraceae, Diegdendraceae, Cchlspermaceae, Peganaceae, Tetradiclidaceae, Nyssaceae, Ternstremiaceae, Pellicieraceae, Aucubaceae, Dnatiaceae, Lbeliaceae, Desfntainiaceae, Diervillaceae, Dipsacaceae, Linnaeaceae, Mrinaceae, and Valerianaceae. The ther discntinued families were: 20

21 Limncharitaceae, Luzuriagaceae, Sparganiaceae, Ledcarpaceae, Heterpyxidaceae, Psilxylaceae, Oliniaceae, Rhynchcalycaceae, Parnassiaceae, Maesaceae, Myrsinaceae, Thephrastaceae, Eremsynaceae, Plysmaceae, Tribelaceae, Aralidiaceae, Mackinlayaceae, and Melanphyllaceae. 20 families were accepted in the APG III system which had nt been in the previus system, and a few families were mved t a different psitin. The newly recgnized families are: Petermanniaceae, Schepfiaceae, Limeaceae, Lphicarpaceae, Mntiaceae, Talinaceae, Anacampsertaceae, Centrplacaceae, Calphyllaceae, Guamatelaceae, Gerrardinaceae, Dipentdntaceae, Capparidaceae, Clemaceae, Cytinaceae, Mitrastemnaceae, Metteniusaceae, Linderniaceae, Thmandersiaceae, and Quintiniaceae. The number f families nt placed in any rder was reduced frm 39 t 10. Apdanthaceae and Cynmriaceae were placed amng the angisperms, incertae sedis, that is, nt in any grup within the angisperms. Eight ther families were placed incertae sedis in varius supra-rdinal grups within the angisperms. The unplaced families were: Apdanthaceae, Cynmriaceae, Dasypgnaceae, Sabiaceae, Dilleniaceae, Icacinaceae, Metteniusaceae, Oncthecaceae, Vahliaceae, and Braginaceae. The circumscriptin f the family Icacinaceae remains especially dubtful. Apdytes, Emmtum, Cassinpsis, and a few ther genera were prvisinally retained within it until further studies can determine whether they prperly belng there. Three genera (Gumillea, Nicbaridendrn, and Petenaea) were placed within the angisperms incertae sedis. Gumillea had been unplaced in APG II. Nicbaridendrn and Petenaea were newly added t the list. The classificatin is shwn belw in tw versins. The shrt versin ges t the level f rders and f families unplaced in an rder. The detailed versin shws all the families. Orders at the same level in the classificatin are arranged alphabetically. Nte that rders may nt cntain the same families as in earlier versins f the APG system (APG system, APG II system). Further detail n relatinships can be seen in the phylgenetic tree belw. Shrt versin clade angisperms rder Ambrellales rder Nymphaeales rder Austrbaileyales rder Chlranthales clade magnliids rder Canellales rder Laurales rder Magnliales rder Piperales clade mncts rder Acrales rder Alismatales rder Asparagales rder Discreales rder Liliales rder Pandanales rder Petrsaviales clade cmmelinids family Dasypgnaceae -- unplaced in an rder rder Arecales rder Cmmelinales rder Pales 21

22 rder Zingiberales prbable sister f eudicts rder Ceratphyllales clade eudicts family Sabiaceae -- unplaced in an rder rder Buxales rder Prteales rder Ranunculales rder Trchdendrales clade cre eudicts family Dilleniaceae -- unplaced in an rder rder Gunnerales rder Saxifragales clade rsids rder Vitales clade fabids (eursids I) rder Celastrales rder Cucurbitales rder Fabales rder Fagales rder Malpighiales rder Oxalidales rder Rsales rder Zygphyllales clade malvids (eursids II) rder Brassicales rder Crsssmatales rder Geraniales rder Huerteales rder Malvales rder Myrtales rder Picramniales rder Sapindales (back t cre eudicts) rder Berberidpsidales rder Caryphyllales rder Santalales clade asterids rder Crnales rder Ericales clade lamiids (euasterids I) family Braginaceae -- unplaced in an rder family Vahliaceae -- unplaced in an rder family Icacinaceae -- unplaced in an rder family Metteniusaceae -- unplaced in an rder family Oncthecaceae -- unplaced in an rder rder Garryales rder Gentianales rder Lamiales rder Slanales 22

23 clade campanulids (euasterids II) rder Apiales rder Aquifliales rder Asterales rder Bruniales rder Dipsacales rder Escallniales rder Paracryphiales Detailed versin (with all families) Legend: * = new family placement; = newly recgnized rder fr the APG system; = new family circumscriptin described in the text; $ = families that represent the brader circumscriptin f ptins available in APG II and favured here; $$ = families that were in square brackets in APG II, the narrwer circumscriptins favured here. Angisperms Ambrellales Melikyan, A.V.Bbrv & Zaytzeva Ambrellaceae Pichn Nymphaeales Salisb. ex Bercht. & J.Presl $$Cabmbaceae Rich. ex A.Rich. *Hydatellaceae U.Hamann $$Nymphaeaceae Salisb. Austrbaileyales Takht. ex Reveal Austrbaileyaceae Crizat $Schisandraceae Blume (including Illiciaceae A.C.Sm.) Trimeniaceae L.S.Gibbs Chlranthales R.Br. Chlranthaceae R.Br. ex Sims Magnliids Canellales Crnquist Canellaceae Mart. Winteraceae R.Br. ex Lindl. Piperales Bercht. & J.Presl Aristlchiaceae Juss. Hydnraceae C.Agardh Lactridaceae Engl. Piperaceae Giseke Saururaceae F.Vigt Laurales Juss. ex Bercht. & J.Presl Atherspermataceae R.Br. Calycanthaceae Lindl. Gmrtegaceae Reiche Hernandiaceae Blume Lauraceae Juss. Mnimiaceae Juss. Siparunaceae Schdde Magnliales Juss. ex Bercht. & J.Presl Annnaceae Juss. 23

24 Degeneriaceae I.W.Bailey & A.C.Sm. Eupmatiaceae Orb. Himantandraceae Diels Magnliaceae Juss. Myristicaceae R.Br. Mncts Acrales Link Acraceae Martinv Alismatales R.Br. ex Bercht. & J.Presl Alismataceae Vent. (including Limncharitaceae Takht. ex Crnquist) Apngetnaceae Planch. Araceae Juss. Butmaceae Mirb. Cymdceaceae Vines Hydrcharitaceae Juss. Juncaginaceae Rich. Psidniaceae Vines Ptamgetnaceae Bercht. & J.Presl Ruppiaceae Hran. Scheuchzeriaceae F.Rudlphi Tfieldiaceae Takht. Zsteraceae Dumrt. Petrsaviales Takht. Petrsaviaceae Hutch. Discreales R.Br. Burmanniaceae Blume Discreaceae R.Br. Nartheciaceae Fr. ex Bjurzn Pandanales R.Br. ex Bercht. & J.Presl Cyclanthaceae Pit. ex A.Rich. Pandanaceae R.Br. Stemnaceae Caruel Triuridaceae Gardner Vellziaceae J.Agardh Liliales Perleb Alstremeriaceae Dumrt. (including Luzuriagaceae Ltsy) Campynemataceae Dumrt. Clchicaceae DC. Crsiaceae Becc. Liliaceae Juss. Melanthiaceae Batsch ex Brkh. *Petermanniaceae Hutch. Philesiaceae Dumrt. Ripgnaceae Cnran & Cliffrd Smilacaceae Vent. Asparagales Link $Amaryllidaceae J.St.-Hil. (including Agapanthaceae F.Vigt, Alliaceae Brkh.) $Asparagaceae Juss. (including Agavaceae Dumrt., Aphyllanthaceae Burnett, Hespercallidaceae Traub, Hyacinthaceae Batsch ex Brkh., Laxmanniaceae Bubani, Ruscaceae M.Rem., Themidaceae Salisb.) 24

25 Asteliaceae Dumrt. Blandfrdiaceae R.Dahlgren & Cliffrd Bryaceae M.W.Chase, Rudall & Cnran Dryanthaceae R.Dahlgren & Cliffrd Hypxidaceae R.Br. Iridaceae Juss. Ixiliriaceae Nakai Lanariaceae R.Dahlgren & A.E.van Wyk Orchidaceae Juss. Tecphilaeaceae Leyb. $Xanthrrheaceae Dumrt. (including Asphdelaceae Juss. and Hemercallidaceae R.Br.) Xernemataceae M.W.Chase, Rudall & M.F.Fay Cmmelinids Dasypgnaceae Dumrt. Arecales Brmhead Arecaceae Bercht. & J.Presl Cmmelinales Mirb. ex Bercht. & J.Presl Cmmelinaceae Mirb. Haemdraceae R. Br. Hanguanaceae Airy Shaw Philydraceae Link Pntederiaceae Kunth Pales Small Anarthriaceae D.F.Cutler & Airy Shaw Brmeliaceae Juss. Centrlepidaceae Endl. Cyperaceae Juss. Ecdeicleaceae D.F.Cutler & Airy Shaw Ercaulaceae Martin v Flagellariaceae Dum rt. Jinvilleaceae Tml. & A.C.Sm. Juncaceae Juss. Mayacaceae Kunth Paceae Barnhart Rapateaceae Dumrt. Restinaceae R.Br. Thurniaceae Engl. Typhaceae Juss. (including Sparganiaceae Hanin) Xyridaceae C.Agardh Zingiberales Griseb. Cannaceae Juss. Cstaceae Nakai Helicniaceae Vines Lwiaceae Ridl. Marantaceae R.Br. Musaceae Juss. Strelitziaceae Hutch. Zingiberaceae Martinv Prbable sister f eudicts 25

26 Ceratphyllales Link Ceratphyllaceae Gray Eudicts Ranunculales Juss. ex Bercht. & J.Presl Berberidaceae Juss. $Circaeasteraceae Hutch. (including Kingdniaceae Airy Shaw) Eupteleaceae K.Wilh. Lardizabalaceae R.Br. Menispermaceae Juss. $Papaveraceae Juss. (including Fumariaceae Marquis, Pteridphyllaceae Nakai ex Reveal & Hgland) Ranunculaceae Juss. Sabiaceae Blume Prteales Juss. ex Bercht. & J.Presl Nelumbnaceae A.Rich. $$Platanaceae T.Lestib. $$Prteaceae Juss. Trchdendrales Takht. ex Crnquist $Trchdendraceae Eichler (including Tetracentraceae A.C.Sm.) Buxales Takht. ex Reveal $Buxaceae Dumrt. (including Didymelaceae Leandri) Cre eudicts Gunnerales Takht. ex Reveal $$Gunneraceae Meisn. $$Myrthamnaceae Nied. Dilleniaceae Salisb. Saxifragales Bercht. & J.Presl Altingiaceae Hran. Aphanpetalaceae Dweld Cercidiphyllaceae Engl. Crassulaceae J.St.-Hil. Daphniphyllaceae Müll.-Arg. Grssulariaceae DC. $$Halragaceae R.Br. Hamamelidaceae R.Br. $Iteaceae J.Agardh (including Pterstemnaceae Small) Paeniaceae Raf. $$Penthraceae Rydb. ex Britt. * Peridiscaceae Kuhlm. (including Medusandraceae Brenan, Syauxia Oliver) Saxifragaceae Juss. $$Tetracarpaeaceae Nakai Berberidpsidales Dweld Aextxicaceae Engl. & Gilg Berberidpsidaceae Takht. Santalales R.Br. ex Bercht. & J.Presl *Balanphraceae Rich. Lranthaceae Juss. Misdendraceae J.Agardh Santalaceae R.Br. 26

27 Olacaceae R.Br. Opiliaceae Valetn *Schepfiaceae Blume Caryphyllales Juss. ex Bercht. & J.Presl Achatcarpaceae Heimerl Aizaceae Martinv Amaranthaceae Juss. *Anacampsertaceae Eggli & Nyffeler Ancistrcladaceae Planch. ex Walp. Asterpeiaceae Takht. ex Reveal & Hgland Barbeuiaceae Nakai Basellaceae Raf. Cactaceae Juss. Caryphyllaceae Juss. Didiereaceae Radlk. Dincphyllaceae Airy Shaw Drseraceae Salisb. Drsphyllaceae Chrtek, Slavíkvá & Studnicka Frankeniaceae Desv. Gisekiaceae Nakai Halphytaceae A.Srian *Limeaceae Shipunv ex Reveal *Lphicarpaceae Dweld & Reveal Mlluginaceae Bartl. *Mntiaceae Raf. Nepenthaceae Dumrt. Nyctaginaceae Juss. Physenaceae Takht. Phytlaccaceae R.Br. Plumbaginaceae Juss. Plygnaceae Juss. Prtulacaceae Juss. Rhabddendraceae Prance Sarcbataceae Behnke Simmndsiaceae Tiegh. Stegnspermataceae Nakai *Talinaceae Dweld Tamaricaceae Link Rsids Vitales Juss. ex Bercht. & J.Presl Vitaceae Juss. Eursids I Zygphyllales Link $$Krameriaceae Dumrt. $$Zygphyllaceae R.Br. Celastrales Link $Celastraceae R.Br. (including Lepurpetalaceae Nakai, Parnassiaceae Martinv, Pttingeriaceae Takht.) Lepidbtryaceae J.Lénard 27

28 Oxalidales Bercht. & J.Presl Brunelliaceae Engl. Cephaltaceae Dumrt. Cnnaraceae R.Br. Cunniaceae R.Br. Elaecarpaceae Juss. ex DC. *Huaceae A.Chev. Oxalidaceae R.Br. Malpighiales Juss. ex Bercht. & J.Presl Achariaceae Harms Balanpaceae Benth. & Hk.f. Bnnetiaceae L.Beauvis. ex Nakai *Calphyllaceae J.Agardh Carycaraceae Vigt *Centrplacaceae Dweld & Reveal $$Chrysbalanaceae R.Br. Clusiaceae Lindl. Ctenlphnaceae Exell & Mendnça $$Dichapetalaceae Baill. Elatinaceae Dumrt. $$ Erythrxylaceae Kunth (including Aneulphus Benth.) Euphrbiaceae Juss. $$Euphrniaceae Marc.-Berti Gupiaceae Miers Humiriaceae A.Juss. Hypericaceae Juss. Irvingiaceae Exell & Mendnça Ixnanthaceae Planch. ex Miq. Lacistemataceae Mart. Linaceae DC. ex Perleb Lphpyxidaceae H. Pfeiff. Malpighiaceae Juss. $Ochnaceae DC. (including Medusagynaceae Engl. & Gilg, Quiinaceae Chisy) Pandaceae Engl. & Gilg $Passiflraceae Juss. ex Russel (including Malesherbiaceae D. Dn, Turneraceae Kunth ex DC.) Phyllanthaceae Martinv Picrdendraceae Small Pdstemaceae Rich. ex Kunth Putranjivaceae Meisn. *Rafflesiaceae Dumrt. $$Rhizphraceae Pers. Salicaceae Mirb. $$Trigniaceae A.Juss. Vilaceae Batsch Cucurbitales Juss. ex Bercht. & J.Presl Anisphylleaceae Ridl. Begniaceae C.Agardh Criariaceae DC. Cryncarpaceae Engl. Cucurbitaceae Juss. 28

29 Datiscaceae Dumrt. Tetramelaceae Airy Shaw Fabales Brmhead Fabaceae Lindl. Plygalaceae Hffmanns. & Link Quillajaceae D.Dn Surianaceae Arn. Fagales Engl. Betulaceae Gray Casuarinaceae R.Br. Fagaceae Dumrt. Juglandaceae DC. ex Perleb (including Rhipteleaceae Hand.-Mazz.) Myricaceae A.Rich. ex Kunth Nthfagaceae Kuprian Ticdendraceae Gómez-Laur. & L.D.Gómez Rsales Bercht. & J.Presl Barbeyaceae Rendle Cannabaceae Martinv Dirachmaceae Hutch. Elaeagnaceae Juss. Mraceae Gaudich. Rhamnaceae Juss. Rsaceae Juss. Ulmaceae Mirb. Urticaceae Juss. Eursids II Geraniales Juss. ex Bercht. & J.Presl $Geraniaceae Juss. (including Hypsecharitaceae Wedd.) $Melianthaceae Hran. (including Francaceae A.Juss.) Vivianiaceae Kltzsch (including Ledcarpaceae Meyen) Myrtales Juss. ex Bercht. & J.Presl Alzateaceae S.A.Graham Cmbretaceae R.Br. Crypterniaceae A.DC. Lythraceae J.St.-Hil. $Melastmataceae Juss. (including Memecylaceae DC.) Myrtaceae Juss. (including Heterpyxidaceae Engl. & Gilg, Psilxylaceae Crizat) Onagraceae Juss. Penaeaceae Sweet ex Guill. (including Oliniaceae Arn., Rhynchcalycaceae L.A.S.Jhnsn & B.G.Briggs) Vchysiaceae A.St.-Hil. Crsssmatales Takht. ex Reveal *Aphliaceae Takht. Crsssmataceae Engl. *Geisslmataceae A.DC. *Guamatelaceae S.Oh & D.Ptter Stachyuraceae J.Agardh Staphyleaceae Martinv * Strasburgeriaceae Sler. (including Ixerbaceae Griseb. ex Dweld & Reveal) Picramniales Dweld 29

30 *Picramniaceae Fernand & Quinn Huerteales Dweld *Dipentdntaceae Merr. *Gerrardinaceae Alfrd Tapisciaceae Takht. Brassicales Brmhead $Akaniaceae Stapf (including Bretschneideraceae Engl. & Gilg) Bataceae Mart. ex Perleb Brassicaceae Burnett *Capparaceae Juss. Caricaceae Dumrt. *Clemaceae Bercht. & J.Presl Emblingiaceae J.Agardh Gyrstemnaceae A.Juss. Keberliniaceae Engl. Limnanthaceae R.Br. Mringaceae Martinv Pentadiplandraceae Hutch. & Dalziel Resedaceae Martinv Salvadraceae Lindl. Setchellanthaceae Iltis Tvariaceae Pax Trpaelaceae Juss. ex DC. Malvales Juss. ex Bercht. & J.Presl $Bixaceae Kunth (including Cchlspermaceae Planch., Diegdendraceae Capurn) Cistaceae Juss. *Cytinaceae A.Rich. Diptercarpaceae Blume Malvaceae Juss. Muntingiaceae C.Bayer, M.W.Chase & M.F.Fay Neuradaceae Kstel. Sarclaenaceae Caruel Sphaersepalaceae Tiegh. ex Bullck Thymelaeaceae Juss. Sapindales Juss. ex Bercht. & J.Presl Anacardiaceae R.Br. Biebersteiniaceae Schnizl. Burseraceae Kunth Kirkiaceae Takht. Meliaceae Juss. $Nitrariaceae Lindl. (including Peganaceae Tiegh. ex Takht., Tetradiclidaceae Takht.) Rutaceae Juss. Sapindaceae Juss. Simarubaceae DC. Asterids Crnales Link. Crnaceae Bercht. & J.Presl (including Nyssaceae Juss. ex Dumrt.) Curtisiaceae Takht. Grubbiaceae Endl. ex Meisn. 30

31 Hydrangeaceae Dumrt. Hydrstachyaceae Engl. Lasaceae Juss. Ericales Bercht. & J.Presl Actinidiaceae Engl. & Gilg. Balsaminaceae A.Rich. Clethraceae Kltzsch Cyrillaceae Lindl. Diapensiaceae Lindl. Ebenaceae Gürke Ericaceae Juss. Fuquieriaceae DC. Lecythidaceae A.Rich. Marcgraviaceae Bercht. & J.Presl *Mitrastemnaceae Makin $Pentaphylacaceae Engl. (including Ternstremiaceae Mirb. ex DC.) Plemniaceae Juss. Primulaceae Batsch ex Brkh. (including Maesaceae Anderb., B.Ståhl & Källersjö, Myrsinaceae R.Br., Thephrastaceae G.Dn) Rridulaceae Martinv Saptaceae Juss. Sarraceniaceae Dumrt. $$Sladeniaceae Airy Shaw Styracaceae DC. & Spreng. Symplcaceae Desf. $Tetrameristaceae Hutch. (including Pellicieraceae L.Beauvis.) Theaceae Mirb. ex Ker Gawl. Euasterids I *Braginaceae Juss. (including Hplestigmataceae Gilg) Vahliaceae Dandy Icacinaceae Miers Metteniusaceae H.Karst. ex Schnizl. Oncthecaceae Kbuski ex Airy Shaw Garryales Lindl. Eucmmiaceae Engl. $Garryaceae Lindl. (including Aucubaceae Bercht. & J.Presl) Gentianales Juss. ex Bercht. & J.Presl Apcynaceae Juss. Gelsemiaceae Struwe & V.A.Albert Gentianaceae Juss. Lganiaceae R.Br. ex Mart. Rubiaceae Juss. Lamiales Brmhead Acanthaceae Juss. Bignniaceae Juss. Byblidaceae Dmin Calcelariaceae Olmstead Carlemanniaceae Airy Shaw Gesneriaceae Rich. & Juss. Lamiaceae Martinv 31

32 *Linderniaceae Brsch, K.Müll., & Eb.Fisch. Lentibulariaceae Rich. Martyniaceae Hran. Oleaceae Hffmanns. & Link Orbanchaceae Vent. Paulwniaceae Nakai Pedaliaceae R.Br. Phrymaceae Schauer Plantaginaceae Juss. Plcspermataceae Hutch. Schlegeliaceae Reveal Scrphulariaceae Juss. Stilbaceae Kunth Tetrachndraceae Wettst. *Thmandersiaceae Sreem. Verbenaceae J.St.-Hil. Slanales Juss. ex Bercht. & J.Presl Cnvlvulaceae Juss. Hydrleaceae R.Br. ex Edwards Mntiniaceae Nakai Slanaceae Juss. Sphencleaceae T.Baskerv. Euasterids II Aquifliales Senft Aquifliaceae Bercht. & J.Presl Cardipteridaceae Blume (including Leptaulaceae Tiegh.) Helwingiaceae Decne. Phyllnmaceae Small Stemnuraceae Kårehed Asterales Link Alseusmiaceae Airy Shaw Argphyllaceae Takht. Asteraceae Bercht. & J.Presl Calyceraceae R.Br. ex Rich. $Campanulaceae Juss. (including Lbeliaceae Juss.) Gdeniaceae R.Br. Menyanthaceae Dumrt. Pentaphragmataceae J.Agardh Phellinaceae Takht. Russeaceae DC. $Stylidiaceae R.Br. (including Dnatiaceae B.Chandler) Escallniales R.Br. Escallniaceae R.Br. ex Dumrt. (including Eremsynaceae Dandy, Plysmaceae Blume, Tribelaceae Airy Shaw) Bruniales Dumrt. Bruniaceae R.Br. ex DC. Clumelliaceae D.Dn (including Desfntainiaceae Endl.) Paracryphiales Takht. ex Reveal Paracryphiaceae Airy Shaw (including *Quintiniaceae Dweld, Sphenstemnaceae P.Ryen & Airy Shaw) 32

33 Dipsacales Juss. ex Bercht. & J.Presl Adxaceae E.Mey. Caprifliaceae Juss. (including Diervillaceae Pyck, Dipsacaceae Juss., Linnaeaceae Backlund, Mrinaceae Raf., Valerianaceae Batsch) Apiales Nakai Apiaceae Lindl. Araliaceae Juss. Griseliniaceae J.R.Frst. & G.Frst. ex A.Cunn. Mydcarpaceae Dweld Pennantiaceae J.Agardh Pittspraceae R.Br. Trricelliaceae Hu (including Aralidiaceae Philipsn & B.C.Stne, Melanphyllaceae Takht. ex Airy Shaw) Taxa f uncertain psitin Apdanthaceae Takht. (three genera) Cynmriaceae Endl. ex Lindl. Gumillea Ruiz & Pav. Petenaea Lundell (prbably in Malvales) Nicbaridendrn (see Simmns, 2004; prbably in Celastraceae). Subfamilies replacing discntinued families The number f Subfamilies replacing APG II bracketed families is reprted in table 1. Table 1 - A number f subfamilies have been prpsed t replace sme f the families which were ptinal (i.e. bracketed) in APG II, but have been discntinued in APG III. [5] These are shwn in the table belw. APG II bracketed family Agapanthaceae Agavaceae Alliaceae Amaryllidaceae Aphyllanthaceae Asparagaceae Asphdelaceae Hemercallidaceae Hespercallidaceae Hyacinthaceae Laxmanniaceae Ruscaceae Themidaceae Xanthrrheaceae Subfamilies replacing APG II bracketed families APG III family: subfamily Amaryllidaceae: Agapanthideae Asparagaceae: Agavideae Amaryllidaceae: Alliideae Amaryllidaceae: Amaryllidideae Asparagaceae: Aphyllanthideae Asparagaceae: Asparagideae Xanthrrheaceae: Asphdelideae Xanthrrheaceae: Hemercallidideae Asparagaceae: Agavideae Asparagaceae: Scillideae Asparagaceae: Lmandrideae Asparagaceae: Nlinideae Asparagaceae: Brdiaeideae Xanthrrheaceae: Xanthrrheideae 33

34 References Angisperm Phylgeny Grup (2009), "An update f the Angisperm Phylgeny Grup classificatin fr the rders and families f flwering plants: APG III", Btanical Jurnal f the Linnean Sciety 161 (2): As easy as APG III - Scientists revise the system f classifying flwering plants, The Linnean Sciety f Lndn, Chase, Mark W. & Reveal, James L. (2009), "A phylgenetic classificatin f the land plants t accmpany APG III", Btanical Jurnal f the Linnean Sciety 161 (2): Chase, M.W.; Reveal, J.L. & Fay, M.F. (2009), "A subfamilial classificatin fr the expanded asparagalean families Amaryllidaceae, Asparagaceae and Xanthrrheaceae", Btanical Jurnal f the Linnean Sciety 161 (2):