Pelargonium
I am attempting to synthesize my observations on molecular phylogenies with the formal classification scheme. Therefore the scheme presented is based heavily on the work of morphological systematists, with annotations for where taxa go in my trees -- Clade 1 is the one containing P. hortorum, Clade 2 is the P. candicans clade, Clade 3 is the P. cotyledonis clade, Clade 4 is the P. capitatum clade. The basic division in the genus by plastid sequences is between Clades 1+2 and 3+4.
Struck 1997 (Pl.Syst.Evol.208:71) presents a scheme based on "karyological and molecular" evidence (from van der Walt and coworkers) and consistent with nectar sugar compositional differences, which breaks the genus into two main clades, as below. Struck's scheme is largely consistent with my analyses and the published molecular phylogenies of Freek Bakker and colleagues (esp see Bakker 2000 Am.J.Bot. 87:727).
- Group A (chromosomes relatively small, putative basic chromosome number 1n=11: Struck 1997; Bakker referred to this as "subgenus Pelargonium" at IBC 1999; primarily from the winter rainfall region/western Cape and exhibiting considerable pollinator dependence)(my structure here reflects the trnLF phylogeny reported in Bakker(1999)PlantSystEvol 216:309, but keeps some sections intact for historical consistency)
- Group A1, for lack of a better term...my two sub-groups within Struck's Group A are based on my ptDNA clades and the data of Bakker 1999 Pl.Syst.Evol 216:309
- section Pelargonium (1n=11; functionally and perhaps evolutionarily most primitive flower morphs in genus: Struck 1997)
- P. betulinum (Struck 1997)
- P. capitatum (Struck 1997) (Clade 4)
- P. crispum (Struck 1997)
- P. cucullatum (Struck 1997)
- P. denticulatum (Struck 1997)
- P. glutinosum (Struck 1997)
- P. papilionaceum (Struck 1997)
- P. scabrum (Struck 1997) (=scabroide?; Struck 1997)
- P. tomentosum (Struck 1997)
- P. setulosum (Clade 4; Mary Gibby places it in this section IBC 1999, says ~50% of individuals are polyploids)
- P. cordatum (synonymous with Pelargonium cordifolium?; here by Bakker 1999)
- P. quercifolium (Bakker 1999)
- P. nanum (Bakker 2000)
- (former) section Glaucophyllum (1n=11) (placement within section Pelargonium by Bakker(1999)PlantSystEvol)
- P. fruticosum (Struck 1997)
- P. laevigatum (Struck 1997)
- P. patulum (Struck 1997)
- P. grandiflorum here? (Clade 4)
- section Campylia (1n=10)
- P. burgerianum (Struck 1997)
- P. coronopifolium (Struck 1997)
- P. ocellatum (Struck 1997)
- P. tricolor (Struck 1997; syn with P. violareum)
- P. incarnatum (Struck 1997)
- P. caespitosum (Struck 1997)
- P. ovale (Struck 1997)
- P. oenothera (Struck 1997)
- P. capillare (Struck 1997)
- P. elegans (Pelargoniums of S.Afr, vol III)
- section Otidia (1n=11)
- P. alternans (Struck 1997) (Clade 4)
- P. crithmifolium (Struck 1997)
- P. ceratophyllum here? (Clade 4)
- section Polyactium (Maggs 1995 S.Afr.J.Bot.61:53; 1n=11; exclusively night-flowering? Struck 1997 p.83)
- subsect Caulescentia (Maggs 1995 S.Afr.J.Bot. 61:173)(Bakker 1998 Pl.Syst.Evol 211:273 places this (by ITS and trnL-F) in Peristera
- subsect Polyactium (putatively primitive: Maggs as cited by Struck 1997)(treated in Maggs 1999 S.Afr.J.Bot. 65:115)(primarily winter rainfall species with underground tuber, pale yellow petals, and very dark purple floral markings; dusk-scented)
- P. pulverulentum (range of ploidy levels)
- P. anethifolium
- P. lobatum
- P. multiradiatum
- P. pillansii
- P. radulifolium
- P. triste (polyploid)
- subsect Magnistipulacea
- P. luridum (actually multiple species? Maggs 1995)
- P. flabellifolium (one of the P. luridum complex; Struck 1997)
- subsect Schizopetala
- P. amatymbicum
- P. bowkeri (Clade 4)
- P. caffrum
- P. schizopetalum
- P. woodii
- P. xerophyton (Knuth 1912 puts this in sect. Ligularia, but Albers 2000 S.Afr.J.Bot. 66:31 does not include it in Ligularia; Bakker 1998 puts it on the outskirts of Peristera, sister to P. gibbosum; my placement is thus somewhat arbitrary -- somewhere in group A1?) (Clade 4)
- section Cortusina (1n=11) (centered in the Northern Cape Province and Namibia: Dreyer 2000)
- P. cortusifolium (Struck 1997)
- P. crassicaule (Struck 1997) (Clade 4)
- P. echinatum (Struck 1997)
- P. magenteum (Struck 1997)
- P. sibthorprifolium (Dreyer 1992)
- section Hoarea (see Marais 1995 S.Afr.J.Bot.61:90, Marais 1997a S.Afr.J.Bot.63:68, Marais 1997b S.Afr.J.Bot.63:82)(at IBC 1999, Gibby reported that DNA data suggest Hoarea monophyly)
- P. aridicola (sp.nov. Marais 1997b)
- P. pubipetalum (sp.nov. Marais 1997b)
- P. hirtipetalum (sp.nov. Marais 1997b)
- P. githagineum (sp.nov. Marais 1998 S.Afr.J.Bot. 64:308)
- P. angustipetalum (sp.nov. Marais 1999 S.Afr.J.Bot. 65:50)
- P. parvipetalum (sp.nov. Marais 1999 S.Afr.J.Bot. 65:50)
- P. rubiginosum (sp.nov. Marais 1999 S.Afr.J.Bot. 65:50)
- P. quarciticola (sp.nov. Meve&Marais, Meve 2000 S.Afr.J.Bot. 66:96)
- P. reflexipetalum (was pulchellum; Marais 1997a)
- P. bubonifolium (was namaquense + congestum; Marais 1997a)
- P. violiflorum (was included in longifolium; Marais 1997a)
- P. aestivale (Marais 1995)
- P. fissifolium (synonym: P.floribundum; Marais 1995)
- P. petroselenifolium (was pilosum; Marais 1995)
- P. aristatum (formerly subsumed into P. barbatum; Marais 1995)
- P. rapaceum (Struck 1997)
- P. carneum (Struck 1997)
- P. dipetalum (Struck 1997)
- P. heterophyllum (Struck 1997)
- P. incrassatum (Struck 1997)
- P. longifolium (Struck 1997)
- P. longiflorum (Struck 1997)
- P. luteolum (Struck 1997)
- P. pilosellifolium (Struck 1997)
- P. pinnatum (Struck 1997)
- P. trifoliatum (Struck 1997)
- P. moniliforme (Struck 1997)
- P. punctatum (Struck 1997)
- P. curviandrum (Struck 1997)
- P. triandrum (Struck 1997)
- P. oblongatum (Struck 1997)
- P. chelidonium (Struck 1997)
- P. auritum (?)
- section Ligularia (treated in Albers 2000 S.Afr.J.Bot. 66:31, on basis of morphology, palynology, karyology, phenolics, and ptDNA trnL-F)(base 1n=11)(this is in Struck's Group B, but by sequence similarity (my analyses and Bakker 1999/Albers 2000) I am putting it here in group A
- P. crassipes
- P. sericifolium (Struck 1997)
- P. hystrix (Struck 1997)
- P. oreophilum (Struck 1997)
- P. hirtum (Struck 1997)
- P. stipulaceum (Struck 1997)
- P. torulosum (Struck 1997)
- P. pulchellum (Struck 1997)
- P. appendiculatum (in Hoarea by Struck 1997 and earlier treatments, but Bakker 1999 Pl.Syst.Evol and Marais 1999 S.Afr.J.Bot. "65:404"--this ref seems to be wrong -- place it here)
- P. fulgidum (Struck 1997; only confirmed bird-pollinated Pelargonium -- pollinated by the Sun Bird, Nectarinia chalybea: Struck 1997; Clade 4; formerly in section Polyactium, but excluded by Maggs 1995 on basis of it not being night-scented and petal color and shape...)
- Group A2, for lack of a better term...my two sub-groups within Struck's Group A are based on my ptDNA clades and the data of Bakker 1999 Pl.Syst.Evol 216:309
- section Peristera (1n=7,8,9,10,11,19 as quoted in Bakker 1998 Pl.Syst.Evol. 211:273; my delineation here is a conservative interpretation of Bakker 1998 (which was based on nuc ITS and pt trnL-F spacer region sequences))
- P. minimum (Struck 1997)
- P. grossularoides (Struck 1997) (by Bakker 1998, this includes P. hypoleucum and P. acugnaticum; probably distributed from E to SW across the cape and onto Tristan da Cunha)
- P. filicaule
- P. iocastum
- P. columbinum
- P. buysii
- (former) section Isopetalum (1n=11)(proposed by van der Walt and Vorster to be primitive in floral morphology, but placed within Peristera, quite derived, by Bakker; level of divergence in ITS suggests a recent (~5 MYR) spread from S.Afr to St. Helena, consistent with a recent origin of the Australian Pelargoniums as well)
- P. cotyledonis (Struck 1997) (Clade 3)
- the Australians (by sequence divergence reported in Bakker 1998, this seems likely a ~5 MYA dispersal from S. Afr)
- P. australe (Clade 3)
- P. drummondii
- P. littorale
- P. erodioides
- P. inodorum
- P. rodneyanum (by Williams 2000 Biochem.Syst.Ecol.)
- (former) section Reniformia (as noted in Bakker 1998, the association of Reniformia with Peristera is also supported by hybridization and karyological data) (treated in Dreyer 2000 S.Afr.J.Bot. 66:44)
- P. reniforme (Clade 3)
- P. sidoides (improperly sidaefolium: Dreyer 1995 S.Afr.J.Bot.61:325; was split off from P. reniforme by Knuth (1912), so I assume it belongs here)
- P. exstipulaceum (Struck 1997; =exstipulatum, I believe (which is Clade 3))
- P. ionidiflorum (Bakker 1998 and references therein)(Clade 3)
- P. album
- P. odoratissimum may belong here (Dreyer 2000, but see also Williams 2000)
- Group B (Chromosomes relatively large, putative basic chromosome number 1n=9: Struck 1997; Bakker referred to this as subgenus Ciconium at IBC 1999; primarily from the eastern Cape, tropical east Africa, Madagascar, the Arabian peninsula, and Asia Minor)
- Group B1, based on my analyses and Bakker 2000
- section Ciconium
- P. alchemilloides (Struck 1997) (Clade 1)(Gibby, IBC: 1n=8; sister taxon is P. elongatum with 1n=4)
- P. articulatum (Struck 1997) (Clade 1; formerly in sect. Ligularia, but here by Albers 1992 and Bakker 2000)
- P. barklyi (Struck 1997) (Clade 1; formerly in section Polyactium, but excluded by Maggs 1995 who say it "deviates far from the circumspection of the section", which is fine as that section probably belongs in my Clade 4)
- P. elongatum (Struck 1997)
- P. peltatum (Struck 1997) (Clade 1)
- P. aridum (Bob Price reports (email, 4-17-96) that this falls in the hortorum/peltatum clade, and therefore is Clade 1)
- P. multibracteatum (Bob Price reports (email, 4-17-96) that this falls in the hortorum/peltatum clade, and therefore is Clade 1)
- P. acetosum (my placement here by virtue of close affinity to the others in the section; Clade 1)
- P. hortorum (my placement here by virtue of close affinity to the others in the section; hybrid of P. zonale and P. inquinans; Clade 1)
- P. zonale (I'm assuming it beongs here)
- P. inquinans (I'm assuming it belongs here)
- P. acraeum (Pelargoniums of S.Afr, vol III)
- P. tongaense (Pelargoniums of S.Afr, vol III; placement confirmed by Bakker 2000)
- P. transvaalense (Pelargoniums of S.Afr, vol III)
- section Subsucculentia (formerly part of Ligularia; van der Walt 1995 S.Afr.J.Bot.61:331)
- P. karooicum (Clade 1; placement is weak, different characters disagree -- see discussion in Bakker 2000)
- P. grandicalcaratum (Clade 1; identical to otaviense in rbcL; potentially bird-pollinated: Struck 1997)
- P. otaviense (Clade 1; identical to grandicalcaratum in rbcL; potentially bird-pollinated: Struck 1997)
- P. spinosum (Clade 1)
- eurasian relatives (on the basis of these two taxa having chomosome # of 2n=34 rather than 2n=20 as the rest of the section, van der Walt 1995 exclude them from the section, but I'm sticking them here on the basis of rbcL trees anyway)
- P. quercetorum (Clade 1)
- P. endlicherianum (Clade 1)
- P. caylae (Pelargoniums of S.Afr, vol III places this in Ciconium, but by Bakker 2000, it belongs at the base of Group B1; note that in pt but not mito trees, it is sister to P. karooicum)
- group B2, based on my analyses and Bakker 2000
- Based on Bakker 2000, this group can be further split into two...
- Group B2i contains section Jenkinsonia (formerly part of Ligularia; van der Walt 1997 S.Afr.J.Bot.63:4)
- P. griseum (Clade 2)
- P. trifidum (Clade 2)
- P. praemorsum (Clade 2)
- P. antidysentericum (Clade 2)
- P. divisifolium
- P. dolomiticum
- P. plurisectum
- P. redactum
- P. seneciodes
- P. tenuicaule
- P. tragacanthoides
- P. plurisectum (Struck 1997)
- P. boranense here? (Clade 2)
- P. mutans (was in Ciconium, here by Bakker 2000)
- Group B2ii contains two sections...
- section Myrrhidium
- P. candicans (Struck 1997) (Clade 2)
- P. longicaule (Struck 1997)
- P. myrrhifolium (Struck 1997)
- P. caucalifolium (Bakker 2000)
- P. whytei (Bakker 2000)
- section Chorisma (formerly part of Ligularia; Albers 1995 S.Afr.J.Bot.61:339)
- P. exhibens (Clade 2)
- P. worcesterae
- P. mollicomum (Clade 2)
- P. tetragonum (Clade 2)
- species which I haven't placed yet...
- Pelargonium alpinum (Clade 3)
- Pelargonium apiifolium (formerly in section Polyactium, but excluded by Maggs 1995 who conclude that it is "not matched with any natural population, and presumed to be of hybrid origin.")
- Pelargonium burtonae
- Pelargonium caffrum
- Pelargonium carnosum
- Pelargonium desertorum (Clade 4)
- Pelargonium fragrans (Hybrid)
- Pelargonium fumariifolium
- Pelargonium glaucifolium (Hybrid)
- Pelargonium graveolens
- Pelargonium ignescens (Hybrid)
- Pelargonium klinghardtense
- Pelargonium laxum
- Pelargonium lobatum
- Pelargonium melissinum (Hybrid)
- Pelargonium myrrhifolium
- Pelargonium palmatum (ptDNA currently being sequenced (mid-2003) by Bob Jansen's group; we'll soon know where it goes!)
- Pelargonium paniculatum (sister to P. xerophyton in Albers 2000, and therefore possibly in Clade 4?)
- Pelargonium roessingense