Disclaimer: this page represents my attempt to synthesize various sources of information about plant phylogeny. It is not intended to be taken as a "system" of taxonomic or phylogenetic organization, and it certainly should not be taken as the "true" phylogeny. It is merely what I am currently inclined to believe, a convenient scheme upon which to tack other pieces of information. Please let me know if I am missing anything that you feel ought to be on here, or if you find anything that you disagree with or that you know to be wrong. Thanks!
Earthly Life/Eukaryota/Mitochondriate Eukaryotes/Critters with spliceosomal introns/Eukaryote "crown"/Plantae
Plantae
(assuming a single primary endosymbiotic origin of plastids)
- Rhodophyta
- Glaucocystophytes
- Viridiplantae (primary green plastids: JDP 10-13-94)(Plastids with Chlorophylls A&B)...accepting that Land Plants are derived Charophytes makes "Viridiplantae" = "Chlorophytes"...
- Chlorarachniophyte NMs (primary green plastid; linked with Cryptophyte NMs (red) by ss rRNA: Bhattacharya&Medlin. But see Van de Peer (1996) site-to-site rate correction analysis; Ishida&C-S at 1998 SMBE put chlorarchnion NM in Ulvophytes)
- Prasinophyceae (=Micromonadophyta; Bhattacharya 1998 SMBE suggests that this group is paraphyletic, with charophytes (including land plants) arising from within, based on actin phylogeny and intron positions)
- Mantoniella
- Nephroselmis
- Tetraselmis=Platymonas
- Mesostigma (sister to land plants in Bhattacharya's 1998 SMBE actin analysis)
- Ulvophyta
- Codium
- Derbesia
- Bryopsis
- Trentepohlia
- Chlorophyta
- Volvox
- Pandorina
- Gonium
- Fritsciella
- Draparnaldia
- Chlorella
- Oedegonium
- Chlamydomonales
- Charophyta
- Raphidonema
- Klebsormidium
- Nitella
- Charales (Chara)
- Zygnematales (Closterium, Spirogyra)
- Coleochetales (Coleochaete)
- LandPlants (widely accepted to be derived charophytes; invasion of the land 400~500 MYA: Gastony lecture, Raven v4, Apdx.D; land plant spores at ~480 MYA by Kenrick&Crane 1997 Nature) ("Byrophytes" refers to liverworts+hornworts+mosses, but is doubtless paraphyletic, so I'll just ignore it...my basic phylogeny is from Yin-Long Qiu's scheme, 1995, and is based mostly on rbcL, morphology, and genomic architecture characters)
- Liverworts(divergence of thallose liverworts from the rest of the land plant trunk ~450 MYA by Kenrick&Crane 1997 Nature, Goremykin 1997 Pl.Syst.Evol.)
- Thallose liverworts (Marchantia)
- Leafy liverworts (Bazzania)
- All other land plants
- Hornworts
- Mosses
- Vascular Plants (origin ~420 MYA, Kenrick&Crane 1997 Nature) ("Pteridophytes" refers to lycopods+Psilotoids+Equisetoids+ferns, but is doubtless paraphyletic, so I am not using it; note that most relationships here are uncertain -- Lycopods, Ferns, Seed Plants are probably robust, but Equisetum and Psilotum are mobile and relationships between the major groups don't seem to be set...)
- Lycopods
- Lycopodiales (Lycopodites ~370 MYR)
- Selaginales (Lepidodendron ~360-290 MYR)
- Isoetales
- Sphenophyta (horsetails; Equisetum)
- Psilotales (placement uncertain: Gastony lecture, 1-16-95, puts it at base of vascular plants; Yin-Long Qiu puts it here, probably as compromise; Chuck's rbcL tree, 12-13-94, puts it within ferns...Manhart Mol Phylog Evol 3:114, 1994 using rbcL puts Psilotales+Ophioglossales; Bob Jansen Science 255:1697 cpDNA rearrangement data that put it in seed-plant+ferns+Psilotum clade vs bryophytes+lycopods; Manhart Am Fern J 85: 182, 1995 using cp SSU rRNA puts Psilotales+Ophioglossales, but gets polyphyletic ferns or lycopods, depending on tree, and Equisetum moves around...)
- Ferns+Seed Plants
- Ferns
- Eusporangiate Ferns (sporangium develops from groups of sporangial initial cells; no specialized mechanism of dehiscence)
- Marattiales
- Ophioglossales
- Leptosporangiate Ferns (sponrangium develops from single sporangial initial cell; dehiscence due to hygroscopic changes in annular cells)
- Osmundaceae
- Schizaeaceae (Lygodium)
- Marsileaceae + Salviniaceae
- Adiantum
- Cheilanthes
- Polypodium
- Asplenium
- Seed Plants ("Gymnosperms" refers to the non-angiosperm-seed-plants, and is probably paraphyletic. Though Shu-Miaw Chaw's nuclear SSU rRNA analyses (MBE 14:56) suggest not. Kenrick&Crane 1997 Nature put seed plant origin at ~380 MYA; Goremykin 1997 Pl.Syst.Evol. mol clock puts angio-conifer split at 350+/-35 MYA.)
- Gingko
- Cycads (Wolfe PNAS 1989 says cycads split from angios 340 MYR; conservatively, 200-340 MYR (ibid,p.6205))
- Conifers (phylogeny from Bob Price talk, 4-13-93, largely based on rbcL(?))
- Pinaceae (Pinus=pine,Picea=spruce,Tsuga=fir,Pseudotsuga=douglas fir)
- the rest
- the southerners
- Auracariaceae
- Podocarpaceae
- the rest
- Scidopitys
- the rest
- Taxaceae+Cephalotaxaceae
- Taxodiaceae (including Cupressaceae)
- Taxodiaceae
- Cupressaceae (Cypress, Juniperus=Cedar)
- More Taxodiaceae
- Gnetales+Angiosperms (maybe a natural group: see, e.g., disagreement of 16S and rbcL...Yin-Long Qiu 4-13-95 estimated this group to be ~250 MYR)
- Gnetales
- Ephedra
- Gnetum
- Welwitschia
- Angiosperms (resolution of major lineages is, by my reckoning, unclear; note that "dicots" includes most basal angiosperm groups as well as monophyletic eudicots, and is thus a paraphyletic group. More realistic character may be pollen, with basal groups monosulcate and eudicots tricolpate. By Raven, 4th ed, p. 585, monosulcate dates to 125 MYR, tricolpate to 120 MYR. Wolfe, PNAS 1989, mol clock dates mono/dicot split at 200+/-40 MYA; Goremykin 1997 Pl.Syst.Evol mol clock says 160+/-16 MYA.)
- Ceratophyllum (basal by big rbcL tree)
- Piperales+Aristolochiales
- Piperales
- Piperaceae (Piper, Peperomia)
- Aristolochiales
- Aristolochiaceae (Asarum, Arisotolochia)
- Nymphaeales (basal and paraphyletic - including all others - by big atpB tree; YQ 10-96:trans-spleced nad1 i4)
- Nymphaeaceae
- Cabomobaceae
- Illiciales
- Chloranthaceae
- Amborellaceae
- Austrobaileyaceae
- monocots (basic groupings based on the combined rbcL/morphological analysis of Chase et al Monocots: systematics and evolution (Kew) 1995. The authors state clearly that these results should be taken as a temporary fix...)
- Acorus (YQ 10-96: cis-spliced nad1.i4)
- the rest
- Aroids+Alismatids (YQ 10-96: cis-spliced nad1.i4; rbcL, rps4, nuc18S VERY weakly put this at base of monocots)
- Commelinoids+Lilianae (YQ 10-96: trans-spliced nad1.i4)
- Commelinoids
- Lilianae
- Asparagales
- Liliales
- Dioscorales
- Laurales
- Magnoliales
- Magnoliaceae
- Annonaceae (Asimina)
- Winteraceae (Drimys)
- eudicots (tricolpate or polyporate pollen; 120 MYr)
- ranunculids
- Lardizabalaceae
- Berberidaceae
- Ranunculaceae
- Fumariaceae+Papaveraceae
- lower hamamelids+"higher" eudicots
- lower ham
- Nelumbo
- Platanus (sycamore)
- Trochodendrales (Trochodendron, Tetracentron)
- "higher" eudicots (caryophilids may be outgroup to other two; see Chase et al 1993 analysis A vs. B; Yin-Long Qiu 4-01-97 puts rosid/asterid split at ~90 MYr)
- caryophillids
- Viscaceae
- Droseraceae
- Plumbaginaceae
- Polygonaceae (note "collar" of fused stipules above leaf axil)
- Caryophyllaceae+Chenopodiaceae+Amaranthaceae (Beta is Chenopod)
- Portulacaceae (Claytonia perfoliata=Miner's Lettuce)
- Phytolacaceae
- Aizoaceae
- Cactaceae ? (Ferocactus, Opuntia)
- rosids (probably (I+(II+Geran))+III); Yin-Long Qiu guesses 60-80 MYr)
- rosid III
- Saxifragaceae sensu stricto (others in the family s.l. are rosid I, asterid I)
- Crassulaceae (Sedum)
- Paeoniaceae
- Grossulariaceae (probably polyphyletic; Ribes)
- Hammamelidaceae (Hammamelis, Liquidamabar)
- rosid II+Geraniaceae-plus (based on my rbcL analyses)
- Sapindales/Malvales/Capparales group
- Sapindales (resinous)
- Aceraceae (Acer)
- Sapindaceae
- Anacardiaceae
- Rutaceae
- Simaroubaceae (Ailanthus)
- Burseraceae
- Malvales (mucilaginous)
- Malvaceae (Malva, Gossypium)
- Tiliaceae
- Sterculiaceae (Theobroma)
- Bombacaceae (baobab)
- Capparales (sensu Rodman?) (glucosinoloates)
- Caricaceae (Carica papaya)
- Limnanthaceae (Limnanthes, Floerkea)
- Akaniaceae
- Tropaeolaceae (Tropaeolum)
- Capparaceae (Cleome, Capparis)
- Brassicaceae
- Myrtales group (placement in Rosid I vs. II unclear)
- Onagraceae (Oenothera, Clarkia)
- Cephalotaceae
- Oxalidaceae (Averrhoa=star friut, Oxalis)
- Geraniaceae-plus (sensu rbcL)
- Crossosoma/Geissoloma/Staphylea group
- Greyia/Francoa/Viviania/Wendtia group
- Geraniaceae (sensu cpDNA)
- Hypseocharis
- Geraniaceae sensu stricto
- rosid I (conservatively, (I5+(I4+I2)+I1+I3). Yin-Long Qiu 9-11-96 says maybe (I5+(I3+(I1+(I2+I4))))...this would give monophyletic group of N-fixing root symbiosis from I-3 up.)
- I-1 (broad Fagales; this grouping based on Manos&Steele (1997) AJB analysis of matK and rbcL sequences. Formerly in the Hamamelididae, and characterized by male catkins, says Julie Mulroy, 4-98, if I got that right)
- Nothofagaceae
- the rest
- Fagaceae (Quercus, Fagus)
- "higher hamamelids"
- Betulaceae (Betula, Alnus)
- Juglandaceae (Carya, Juglans)
- Myricaceae
- Rhoiptelea, Casuarina, Ticodendron
- I-2 ("Cucurbitales")
- Cucurbitaceae
- Begoniaceae
- Datiscaceae
- I-3 (broad Legumes)
- I-4 (broad Urticales)
- Ulmaceae
- Urticaceae
- Moraceae
- Rhamnaceae
- Rosaceae (Julie Mulroy 5-98: Prunus distinguished by glands on stipule and sharp bitter scent from scratched bark)
- I-5
- Malpighiaceae
- Euphorbiaceae
- Salicacee
- Passifloraceae
- Chrysobalanaceae
- Violaceae (Viola)
- Linaceae
- Celastraceae (Celastrus, Euonymous)
- asterids ((((I+II)+III)+IV)+V) (sympetalous, tenuinucellate embryo -- megasporocyte is only one cell layer deep)
- asterid V
- asterid IV
- Cornaceae
- Nyssaceae
- Hydrangaceae
- asterid III
- Polemoniaceae (Phlox, Gaultheria)
- Fouquieriaceae
- Balsaminaceae (Impatiens)
- Dilleniidae, mostly
- Ericaceae (Gilia)
- Theaceae (Camellia)
- Primulaceae
- asterid II
- Asteraceae (Kim&Jansen AJB 1995: 23-43 MYR by molclock)
- Caprifoliaceae (Viburnum, Sambucus)
- Valerianaceae
- Aquifoliaceae (Ilex)
- Apiaceae+Araliaceae (note the way compound leaves attach to stem; Fatsia, Hedera are Araliaceae)
- Campanulaceae+Lobeliaceae (anthers come off of ovary separate, but join (Campanulaceae) or fuse (Lobeliaceae) above ovary, forcing stigma to push up through)
- asterid I
- Solanaceae/Convulvulaceae/Boraginaceae group (~Solanales+)
- Solanaceae
- Convulvulaceae
- Boraginaceae
- Hydrophyllaceae
- Asclepiadaceae+Apocynaceae (Vinca)
- Gentianaceae
- Rubiaceae (Galium, Coffea)
- Loganiaceae
- Mints+Scrophs+friends (do all have anthers fused to petals?)
- Lamiaceae+Verbenaceae
- Lamiaceae have 4-lobed schizocarp;the style arises between them (Yin-Long Qiu, 8-19-96)
- Verbenaceae have entire ovary; the style arises from it (Tektona=teak)
- Scophulariaceae+Plantaginaceae
- Acanthaceae
- Bignoniaceae (Catalpa)
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