Relationships among the perennial, endemic umbellifers of

western NA

Stephen R. Downie

Feng-Jie Sun

Ronald L. Hartman

Meet Ph.D. Student Feng-Jie Sun!A phylogenetic study of Cymopterus and related genera

Meet NA Umbellifer Cymopterus(the spring-parsleys)

Cymopterus montanus(RM Herbarium Specimen)

Cymopterus purpureus

• Perennial, taprooted herbs• Low-growing, acaulescent• Ternately-cleft, compound

or dissected leaves• Spreading compound umbel• Yellow or white flowers,

opening very early in spring

• Mericarps conspicuously winged (kyma=wave; pteron=wing)

Number of species per state:1-4 5-8 9-12 13-15 16-18

BONAP(TAMU)

Cymopterus (35 spp.) is distributed primarily throughout the Intermountain and

Montane regions of western NA

Probable relationship of Cymopterus and its alliesA monograph of Cymopterus including a critical study of related genera (Mathias 1930)

UC TV

Cymopterus is putatively allied to other indigenous, perennial umbellifers of western NA

(number of species according to Kartesz, 1994)

• Aletes (6)• Glehnia (1)• Harbouria (1)• Lomatium (78)• Musineon (4)• Neoparrya (1)• Oreonana (3)

• Oreoxis (4)• Orogenia (2)• Podistera (4)• Pseudocymopterus (2)• Pteryxia (4)• Shoshonea (1)• Tauschia (10)

Lomatium grayi

dorsal ribs

lateral wings

Cymopterus purpureus

wings

Dorsal ribs of the fruit (or at least one of them) generally more or less conspicuously winged, rarely all wingless …………... Cymopterus

Dorsal ribs of the fruit wingless or nearly so ………….. Lomatium

Cronquist (1997), treatment of Apiaceae for Intermountain Flora

Cymopterus lapidosus

lateral wings

dorsal ribs

“the distinction between Cymopterus and Lomatium is subject to failure”

“Both genera are of considerable size, and our taxonomic understanding would not be improved by combining them into one.” Cronquist, 1997, Intermountain Flora

Several species of Cymopterus lack dorsal wings and their fruits show similarities to those of typical

Lomatium

C. corrugatusC. deserticolaC. douglassiiC. lapidosusC. megacephalusC. newberryiC. ripleyiC. williamsii

Cymopterus williamsiiHartman & Constance (1985)

Pseudocymopterus montanus

wings

lateral wings

dorsal ribs

Pseudocymopterus, “one of the most complex situations in the family” (Mathias 1930)

Cronquist, 1997

Short, stiff pubescenceat top of peduncleDorsal wings range from well-developed to low

ridges, or are completely absent

Plants show similarities to Cymopterus & Lomatium

Median cross-sections of mature mericarps of Cymopterus (Mathias, 1930) Figs. 1-14

Fig. 15 Fig. 16Fig. 17Fig. 18Fig. 19Fig. 20Fig. 21Figs. 22-23

C. fendleriC. cinerariusC. globosusC. megacephalusC. deserticolaC. coulteriC. corrugatusC. acaulisC. newberryi

Aletes humilis

AulospermumA. panamintenseA. duchesnenseA. minimusA. roseiA. aboriginum

PteryxiaP. terebinthina var. californica var. foeniculacea var. albiflora var. calcareaP. petraea

PseudocymopterusP. montanusP. davidsoniiP. anisatusP. hendersoniiP. humboldtensis

Median cross-sections of mature mericarps (Mathias, 1930)

Taxonomic Confusion Surrounds Cymopterus!

Mathias & Constance (1944-1945) submerged Aulospermum, Phellopterus and most

Pseudocymopterus into Cymopterus (32 spp.), while maintaining Rhysopterus as

distinct

Coulter & Rose (1900) and Mathias (1930) recognized many small, generically distinct elements (e.g., Aulospermum, Phellopterus, Pseudocymopterus, Rhysopterus)

Jones (1908) recognized multiple sections and subgroups within a highly variable and expanded genus Cymopterus

UCTV

The Regents of the University of California

Cronquist (1961, 1997) included Oreoxis, Pseudocymopterus and Pteryxia within a broadly circumscribed Cymopterus (with ca. 45 species)

Kartesz (1994), whose checklist of NA Apiaceae was influenced by Lincoln Constance, maintained these

three genera as distinct, and recognized 35 species in Cymopterus

Contemporary Treatments Recognize 35-45 Species in Cymopterus

Hunt Institute

The Regents of the University of California

“The taxonomic definition of Cymopterus is a vexed question.”

CymopterusPteryxiaGlehniaOrogeniaPrionosciadiumLomatiumPolytaeniaHarboureaPseudocymopterusAralia1.0 changeLength = 14 stepsCI = 0.7143RI= 0.7143

Phylogram derived from MP analysis of 10 morphological characters (data set reanalyzed to include Lomatium alliance and 3 outgroup taxa)

Relationships of Lomatium among other genera of Apiaceae (Plant Syst. Evol., Gilmartin & Simmons, 1987)

• Attempted to discern sister taxon to Lomatium

• 88 NA genera clustered into 7 phenetic alliances based on development of a stylopodium, presence of a carpophore, and direction of fruit flattening

• Monophyly of all genera assumed

Objectives

• To evaluate the monophyly of Cymopterus

• To determine the phylogenetic relationships of the elements that currently comprise Cymopterus with other perennial, endemic apioid umbellifers of western NA

• To evaluate the monophyly of these perennial, endemic umbellifers and to ascertain its sister group

Relationships among NA ApioideaeMP Strict Consensus Tree Showing Tribes and Major Clades

Aethusa cynapiumBifora radiansCoriandrum sativumHeracleum sphondyliumPastinaca sativaConium maculatumLevisticum officinalePimpinella saxifragaAmmi majusPetroselinum crispumAnethum graveolensFoeniculum vulgareApium graveolensFalcaria vulgarisCarum carviAegopodium podagrariaTrachyspermum copticumAnthriscus caucalisMyrrhis odorataOsmorhiza longistylisScandix pecten-venerisChaerophyllum tainturieriCuminum cyminumDaucus pusillusCaucalis platycarposTurgenia latifoliaTorilis arvensisYabea microcarpaLigusticum scoticumBerula erectaSium suaveHelosciadium nodiflorumOxypolis rigidiorCryptotaenia canadensisCynosciadium digitatumLilaeopsis carolinensisCicuta maculata angustifoliaOenanthe pimpinelloidesPerideridia kelloggiiLigusticum porteriConioselinum scopulorumConioselinum chinenseLigusticum canadenseErigenia bulbosaPhysospermum cornubiensePleurospermum foetensOenantheaeScandiceaeCoriandreaeHeracleum cladeApium cladeCareaeConioselinum cladePleurospermeae959110010069999768529293988875756177545497916670100Angelica & Arracacia clades

• 78 genera (333 spp.) of subfamily Apioideae native or naturalized in NA (north of Mexico; Kartesz, 1994)

• All but 3 genera included in molecular analyses to date

• Approximately 30 NA apioid genera in Angelica and Arracacia clades

Materials and Methods

159 Accessions Examined, including all species and infraspecific taxa of Aletes, Cymopterus, Glehnia, Musineon, Oreonana, Oreoxis, Orogenia, Podistera, Polytaenia, Pseudocymopterus, and Pteryxia. Also, Lomatium (30 accessions and Tauschia (5 accessions), and the monotypic Harbouria, Neoparrya, and Shoshonea.

18S nrDNA26S nrDNAITS 5ITS 3ITS 4ITS 2ITS 2ITS 15.8S215 bp224 bpITS region

As putative outgroups, we examined Angelica, 3 representatives of the meso-American Arracacia clade, and several members of the perennial, circumboreal genera Seseli, Selinum, and Peucedanum. The trees were rooted with Aethusa.

Phylogenetic Analyses:• Maximum Likelihood (PAUP*)• Maximum Parsimony (PAUP*)• Bayesian Inference (MrBayes)

Sequence Characteristic

Length variation (bp)No. of aligned positionsNo. of positions constantNo. of positions informativeNo. of positions autapomorphicNo. of alignment gapsNo. of gaps parsimony informativeMaximum sequence divergence

ITS-1

206-218225918945211112.5%

ITS-2

220-22623393796116513.1%

Combined(ITS-1 & ITS-2)

427-441458184168 (37%)106371612.1%

Multiple alignment of 159 ITS-1 and ITS-2 DNA sequences

Angelica argutaAngelica breweriAngelica grayiAngelica pinnataAngelica roseanaAngelica sylvestrisSphenosciadium capitellatumGlehnia littoralis subsp. leiocarpaAngelica amplaAngelica archangelicaSeseli tortuosumSeseli gracilePeucedanum japonicumGlehnia littoralis subsp. littoralisSeseli libanotisSelinum candolleiRhodosciadium argutumPrionosciadium turneriArracacia brandegeiCiclospermum leptophyllumSpermolepis inermisPeucedanum ostruthiumSelinum carvifoliaAethusa cynapium9562668152719772969584Angelica argutaAngelica breweriAngelica grayiAngelica pinnataAngelica roseanaSphenosciadium capitellatumAngelica amplaAngelica archangelicaAngelica sylvestrisGlehnia littoralis subsp. leiocarpaSeseli tortuosumSeseli gracilePeucedanum japonicumGlehnia littoralis subsp. littoralisSelinum candolleiSeseli libanotisPrionosciadium turneriRhodosciadium argutumArracacia brandegeiCiclospermum leptophyllumSpermolepis inermisPeucedanum ostruthiumSelinum carvifoliaAethusa cynapium9179765889737297605799759994135 accessions(18 genera)135 accessions(18 genera)

Basal Relationships: Monophyly of Perennial, Endemic NA Apioideae

MP Strict Consensus Maximum Likelihood

Br. Alfred BrousseauSaint Mary’s College

Peucedanum japonicum

Br. Alfred BrousseauSt. Mary’s College

Glehnia littoralis subsp. leiocarpa

Angelica hendersonii

Charles Webber CA Academy Sciences

Glehnia, “a single species complex occurring on both shores of the

northern Pacific”

subsp. littoralis

Hiroe, 1962)

subsp. leiocarpa

Mathias, 1928

)Cymopterus globosus (CalPhotos) Pteryxia terebinthina (CalPhotos)

C. multinervatus (Cronquist 1997)

Podistera

Largely western NA endemics(north of Mexico)

Obsolete stylopodium(except in Podistera)

Evidence for monophyly

)

M. Schlessman & F. Barrie• Floral protogyny• Nectary morphology• Biogeography• Habitat• Flowering time• Variation in patterns of sex expression

Perennials

Low growingherbaceous perennials

Xerophytic or semixerophytic habit(occur in dry, sandy or alkaline habitats)

Montane or alpine

Overlapping patterns of similar fruit morphology

Cymopterus basalticusCymopterus gilmaniiCymopterus globosusCymopterus ripleyiLomatium juniperinumLomatium orientaleOrogenia fusiformisOrogenia linearifoliaLomatium dasycarpum subsp. dasycarpumLomatium foeniculaceum subsp. foeniculaceumCymopterus lapidosusCymopterus aboriginumCymopterus cinerariusCymopterus evertiiCymopterus panamintensis var. panamintensisOreonana purpurascensOreonana clementisLomatium cousLomatium macrocarpumOreonana vestitaLomatium grayi var. grayiLomatium grayi var. depauperatumCymopterus glaucusLomatium bradshawiiLomatium nudicauleLomatium bicolor var. bicolorLomatium triternatum subsp. platycarpumLomatium californicumLomatium idahoenseCymopterus longipesCymopterus planosusLomatium ambiguumCymopterus davisiiPteryxia terebinthina var. foeniculaceaPteryxia terebinthina var. terebinthinaPteryxia terebinthina var. californicaLomatium dissectum var. dissectumLomatium rigidumTauschia parishiiLomatium repostumLomatium shevockiiLomatium lucidumTauschia argutaCymopterus duchesnensisCymopterus goodrichiiPodistera nevadensisCymopterus minimusCymopterus purpureusCymopterus roseiCymopterus jonesiiLomatium scabrum var. scabrumLomatium parvifoliumTauschia glaucaTauschia kelloggiiLomatium howelliiCymopterus panamintensis var. acutifoliusShoshonea pulvinataTauschia texanaMusineon vaginatumOreoxis alpina subsp. alpinaLomatium greenmaniiTaenidia integerrimaLomatium latilobumLomatium graveolens var. graveolensPteryxia hendersoniiCymopterus nivalisLomatium junceumAletes humilisAletes acaulisMusineon tenuifoliumOreoxis humilisCymopterus williamsiiAletes anisatusCymopterus longilobusLomatium nuttalliiPteryxia hendersoniiLomatium brandegeiPodistera yukonensisNeoparrya lithophilaPodistera eastwoodiaeMusineon lineareCymopterus acaulis var. fendleriCymopterus acaulis var. acaulisCymopterus acaulis var. higginsiiCymopterus newberryiLomatium concinnumCymopterus acaulis var. greeleyorumCymopterus acaulis var. parvusCymopterus bulbosusPteryxia terebinthina var. albiflora (3)Pteryxia terebinthina var. calcarea (3)Pteryxia petraeaPteryxia terebinthina var. calcareaCymopterus corrugatusCymopterus ibapensisCymopterus coulteriCymopterus deserticolaAletes sessiliflorusOreoxis bakeriAletes filifoliusHarbouria trachypleuraPseudocymopterus longiradiatusAletes macdouglaii subsp. macdouglaiiCymopterus beckiiPseudocymopterus montanusPteryxia davidsoniiThaspium barbinodeThaspium pinnatifidumThaspium trifoliatum var. trifoliatumZizia aureaZizia apteraPolytaenia nuttalliiPolytaenia texanaCymopterus douglassiiMusineon divaricatum var. divaricatumCymopterus macrorhizusCymopterus montanusCymopterus constancei (2)Cymopterus multinervatusCymopterus purpurascensAletes calcicolaAletes macdougalii subsp. breviradiatus (2)Aletes macdougalii subsp. breviradiatus (3)Oreoxis trotteri (2)Podistera macouniiPodistera macounii 565897818152671009464617179818770676384558210056951007873969477 Cymopterus basalticusCymopterus gilmaniiCymopterus globosusCymopterus ripleyiLomatium juniperinumLomatium orientaleOrogenia fusiformisOrogenia linearifoliaLomatium dasycarpum subsp. dasycarpumLomatium foeniculaceum subsp. foeniculaceumCymopterus lapidosusCymopterus aboriginumCymopterus cinerariusCymopterus evertiiCymopterus panamintensis var. panamintensisOreonana purpurascensOreonana clementisLomatium cousLomatium macrocarpumOreonana vestitaLomatium grayi var. grayiLomatium grayi var. depauperatumCymopterus glaucusLomatium bradshawiiLomatium nudicauleLomatium bicolor var. bicolorLomatium triternatum subsp. platycarpumLomatium californicumLomatium idahoenseCymopterus longipesCymopterus planosusLomatium ambiguumCymopterus davisiiPteryxia terebinthina var. foeniculaceaPteryxia terebinthina var. terebinthinaPteryxia terebinthina var. californicaLomatium dissectum var. dissectumLomatium rigidumTauschia parishiiLomatium repostumLomatium shevockiiLomatium lucidumTauschia argutaCymopterus duchesnensisCymopterus goodrichiiPodistera nevadensisCymopterus minimusCymopterus purpureusCymopterus roseiCymopterus jonesiiLomatium scabrum var. scabrumLomatium parvifoliumTauschia glaucaTauschia kelloggiiLomatium howelliiCymopterus panamintensis var. acutifoliusShoshonea pulvinataTauschia texanaMusineon vaginatumOreoxis alpina subsp. alpinaLomatium greenmaniiTaenidia integerrimaLomatium latilobumLomatium graveolens var. graveolensPteryxia hendersoniiCymopterus nivalisLomatium junceumAletes humilisAletes acaulisMusineon tenuifoliumOreoxis humilisCymopterus williamsiiAletes anisatusCymopterus longilobusLomatium nuttalliiPteryxia hendersoniiLomatium brandegeiPodistera yukonensisNeoparrya lithophilaPodistera eastwoodiaeMusineon lineareCymopterus acaulis var. fendleriCymopterus acaulis var. acaulisCymopterus acaulis var. higginsiiCymopterus newberryiLomatium concinnumCymopterus acaulis var. greeleyorumCymopterus acaulis var. parvusCymopterus bulbosusPteryxia terebinthina var. albiflora (3)Pteryxia terebinthina var. calcarea (3)Pteryxia petraeaPteryxia terebinthina var. calcareaCymopterus corrugatusCymopterus ibapensisCymopterus coulteriCymopterus deserticolaAletes sessiliflorusOreoxis bakeriAletes filifoliusHarbouria trachypleuraPseudocymopterus longiradiatusAletes macdouglaii subsp. macdouglaiiCymopterus beckiiPseudocymopterus montanusPteryxia davidsoniiThaspium barbinodeThaspium pinnatifidumThaspium trifoliatum var. trifoliatumZizia aureaZizia apteraPolytaenia nuttalliiPolytaenia texanaCymopterus douglassiiMusineon divaricatum var. divaricatumCymopterus macrorhizusCymopterus montanusCymopterus constancei (2)Cymopterus multinervatusCymopterus purpurascensAletes calcicolaAletes macdougalii subsp. breviradiatus (2)Aletes macdougalii subsp. breviradiatus (3)Oreoxis trotteri (2)Podistera macouniiPodistera macounii 565897818152671009464617179818770676384558210056951007873969477 Cymopterus basalticusCymopterus gilmaniiCymopterus globosusCymopterus ripleyiLomatium juniperinumLomatium orientaleOrogenia fusiformisOrogenia linearifoliaLomatium dasycarpum subsp. dasycarpumLomatium foeniculaceum subsp. foeniculaceumCymopterus lapidosusCymopterus aboriginumCymopterus cinerariusCymopterus evertiiCymopterus panamintensis var. panamintensisOreonana purpurascensOreonana clementisLomatium cousLomatium macrocarpumOreonana vestitaLomatium grayi var. grayiLomatium grayi var. depauperatumCymopterus glaucusLomatium bradshawiiLomatium nudicauleLomatium bicolor var. bicolorLomatium triternatum subsp. platycarpumLomatium californicumLomatium idahoenseCymopterus longipesCymopterus planosusLomatium ambiguumCymopterus davisiiPteryxia terebinthina var. foeniculaceaPteryxia terebinthina var. terebinthinaPteryxia terebinthina var. californicaLomatium dissectum var. dissectumLomatium rigidumTauschia parishiiLomatium repostumLomatium shevockiiLomatium lucidumTauschia argutaCymopterus duchesnensisCymopterus goodrichiiPodistera nevadensisCymopterus minimusCymopterus purpureusCymopterus roseiCymopterus jonesiiLomatium scabrum var. scabrumLomatium parvifoliumTauschia glaucaTauschia kelloggiiLomatium howelliiCymopterus panamintensis var. acutifoliusShoshonea pulvinataTauschia texanaMusineon vaginatumOreoxis alpina subsp. alpinaLomatium greenmaniiTaenidia integerrimaLomatium latilobumLomatium graveolens var. graveolensPteryxia hendersoniiCymopterus nivalisLomatium junceumAletes humilisAletes acaulisMusineon tenuifoliumOreoxis humilisCymopterus williamsiiAletes anisatusCymopterus longilobusLomatium nuttalliiPteryxia hendersoniiLomatium brandegeiPodistera yukonensisNeoparrya lithophilaPodistera eastwoodiaeMusineon lineareCymopterus acaulis var. fendleriCymopterus acaulis var. acaulisCymopterus acaulis var. higginsiiCymopterus newberryiLomatium concinnumCymopterus acaulis var. greeleyorumCymopterus acaulis var. parvusCymopterus bulbosusPteryxia terebinthina var. albiflora (3)Pteryxia terebinthina var. calcarea (3)Pteryxia petraeaPteryxia terebinthina var. calcareaCymopterus corrugatusCymopterus ibapensisCymopterus coulteriCymopterus deserticolaAletes sessiliflorusOreoxis bakeriAletes filifoliusHarbouria trachypleuraPseudocymopterus longiradiatusAletes macdouglaii subsp. macdouglaiiCymopterus beckiiPseudocymopterus montanusPteryxia davidsoniiThaspium barbinodeThaspium pinnatifidumThaspium trifoliatum var. trifoliatumZizia aureaZizia apteraPolytaenia nuttalliiPolytaenia texanaCymopterus douglassiiMusineon divaricatum var. divaricatumCymopterus macrorhizusCymopterus montanusCymopterus constancei (2)Cymopterus multinervatusCymopterus purpurascensAletes calcicolaAletes macdougalii subsp. breviradiatus (2)Aletes macdougalii subsp. breviradiatus (3)Oreoxis trotteri (2)Podistera macouniiPodistera macounii 565897818152671009464617179818770676384558210056951007873969477

Cymopterus, Lomatium, and Pteryxia are each highly polyphyleticML analysis of 135 accessions (18 genera) of NA Apioideae

Cymopterusn=41

Lomatiumn=30

Pteryxian=10

Cymopterus basalticusCymopterus gilmaniiCymopterus globosusCymopterus ripleyiLomatium juniperinumLomatium orientaleOrogenia fusiformisOrogenia linearifoliaLomatium dasycarpum subsp. dasycarpumLomatium foeniculaceum subsp. foeniculaceumCymopterus lapidosusCymopterus aboriginumCymopterus cinerariusCymopterus evertiiCymopterus panamintensis var. panamintensisOreonana purpurascensOreonana clementisLomatium cousLomatium macrocarpumOreonana vestitaLomatium grayi var. grayiLomatium grayi var. depauperatumCymopterus glaucusLomatium bradshawiiLomatium nudicauleLomatium bicolor var. bicolorLomatium triternatum subsp. platycarpumLomatium californicumLomatium idahoenseCymopterus longipesCymopterus planosusLomatium ambiguumCymopterus davisiiPteryxia terebinthina var. foeniculaceaPteryxia terebinthina var. terebinthinaPteryxia terebinthina var. californicaLomatium dissectum var. dissectumLomatium rigidumTauschia parishiiLomatium repostumLomatium shevockiiLomatium lucidumTauschia argutaCymopterus duchesnensisCymopterus goodrichiiPodistera nevadensisCymopterus minimusCymopterus purpureusCymopterus roseiCymopterus jonesiiLomatium scabrum var. scabrumLomatium parvifoliumTauschia glaucaTauschia kelloggiiLomatium howelliiCymopterus panamintensis var. acutifoliusShoshonea pulvinataTauschia texanaMusineon vaginatumOreoxis alpina subsp. alpinaLomatium greenmaniiTaenidia integerrimaLomatium latilobumLomatium graveolens var. graveolensPteryxia hendersoniiCymopterus nivalisLomatium junceumAletes humilisAletes acaulisMusineon tenuifoliumOreoxis humilisCymopterus williamsiiAletes anisatusCymopterus longilobusLomatium nuttalliiPteryxia hendersoniiLomatium brandegeiPodistera yukonensisNeoparrya lithophilaPodistera eastwoodiaeMusineon lineareCymopterus acaulis var. fendleriCymopterus acaulis var. acaulisCymopterus acaulis var. higginsiiCymopterus newberryiLomatium concinnumCymopterus acaulis var. greeleyorumCymopterus acaulis var. parvusCymopterus bulbosusPteryxia terebinthina var. albiflora (3)Pteryxia terebinthina var. calcarea (3)Pteryxia petraeaPteryxia terebinthina var. calcareaCymopterus corrugatusCymopterus ibapensisCymopterus coulteriCymopterus deserticolaAletes sessiliflorusOreoxis bakeriAletes filifoliusHarbouria trachypleuraPseudocymopterus longiradiatusAletes macdouglaii subsp. macdouglaiiCymopterus beckiiPseudocymopterus montanusPteryxia davidsoniiThaspium barbinodeThaspium pinnatifidumThaspium trifoliatumZizia aureaZizia apteraPolytaenia nuttalliiPolytaenia texanaCymopterus douglassiiMusineon divaricatum var. divaricatumCymopterus macrorhizusCymopterus montanusCymopterus constanceiCymopterus multinervatusCymopterus purpurascensAletes calcicolaAletes macdougalii subsp. breviradiatus (2)Aletes macdougalii subsp. breviradiatus (3)Oreoxis trotteri (2)Podistera macouniiPodistera macounii 565897818152671009464617179818770676384558210056951007873969477100 Cymopterus basalticusCymopterus gilmaniiCymopterus globosusCymopterus ripleyiLomatium juniperinumLomatium orientaleOrogenia fusiformisOrogenia linearifoliaLomatium dasycarpum subsp. dasycarpumLomatium foeniculaceum subsp. foeniculaceumCymopterus lapidosusCymopterus aboriginumCymopterus cinerariusCymopterus evertiiCymopterus panamintensis var. panamintensisOreonana purpurascensOreonana clementisLomatium cousLomatium macrocarpumOreonana vestitaLomatium grayi var. grayiLomatium grayi var. depauperatumCymopterus glaucusLomatium bradshawiiLomatium nudicauleLomatium bicolor var. bicolorLomatium triternatum subsp. platycarpumLomatium californicumLomatium idahoenseCymopterus longipesCymopterus planosusLomatium ambiguumCymopterus davisiiPteryxia terebinthina var. foeniculaceaPteryxia terebinthina var. terebinthinaPteryxia terebinthina var. californicaLomatium dissectum var. dissectum56589781

37 taxa in Clade:Cymopterus (13)Lomatium (16)Oreonana (3)Pteryxia (3)Orogenia (2)

Maximum Parsimony,Maximum Likelihood, &Bayesian Trees all reveal similar major clades, but resolution within and among them is poor

Cymopterus and Lomatium are highly polyphyletic, with their species inextricably linked with those of Aletes, Musineon, Oreonana, Oreoxis, Orogenia, Podistera, Pseudocymopterus, Pteryxia, and Tauschia

ML analysis135 accessions(18 genera)

AletesCymopterusLomatiumMusineonOreonanaOreoxisOrogeniaPodisteraPseudocymopterusPteryxiaTauschia

94130434252105

17354573151741419

Genus No. constrainedaccessions

Additional steps required to force monophyly

(unconstrained MP analysis resulted in minimal length trees of 881 steps)

Many western NA apioid genera are not monophyletic

Thaspium barbinodeThaspium pinnatifidumThaspium trifoliatum Zizia aureaZizia apteraPolytaenia nuttalliiPolytaenia texana1005695100100

Thaspium, Zizia and Polytaenia are each monophyletic and unite as a clade

Cymopterus basalticusCymopterus gilmaniiCymopterus globosusCymopterus ripleyiLomatium juniperinumLomatium orientaleOrogenia fusiformisOrogenia linearifoliaLomatium dasycarpum subsp. dasycarpumLomatium foeniculaceum subsp. foeniculaceumCymopterus lapidosusCymopterus aboriginumCymopterus cinerariusCymopterus evertiiCymopterus panamintensis var. panamintensisOreonana purpurascensOreonana clementisLomatium cousLomatium macrocarpumOreonana vestitaLomatium grayi var. grayiLomatium grayi var. depauperatumCymopterus glaucusLomatium bradshawiiLomatium nudicauleLomatium bicolor var. bicolorLomatium triternatum subsp. platycarpumLomatium californicumLomatium idahoenseCymopterus longipesCymopterus planosusLomatium ambiguumCymopterus davisiiPteryxia terebinthina var. foeniculaceaPteryxia terebinthina var. terebinthinaPteryxia terebinthina var. californicaLomatium dissectum var. dissectumLomatium rigidumTauschia parishiiLomatium repostumLomatium shevockiiLomatium lucidumTauschia argutaCymopterus duchesnensisCymopterus goodrichiiPodistera nevadensisCymopterus minimusCymopterus purpureusCymopterus roseiCymopterus jonesiiLomatium scabrum var. scabrumLomatium parvifoliumTauschia glaucaTauschia kelloggiiLomatium howelliiCymopterus panamintensis var. acutifoliusShoshonea pulvinataTauschia texanaMusineon vaginatumOreoxis alpina subsp. alpinaLomatium greenmaniiTaenidia integerrimaLomatium latilobumLomatium graveolens var. graveolensPteryxia hendersoniiCymopterus nivalisLomatium junceumAletes humilisAletes acaulisMusineon tenuifoliumOreoxis humilisCymopterus williamsiiAletes anisatusCymopterus longilobusLomatium nuttalliiPteryxia hendersoniiLomatium brandegeiPodistera yukonensisNeoparrya lithophilaPodistera eastwoodiaeMusineon lineareCymopterus acaulis var. fendleriCymopterus acaulis var. acaulisCymopterus acaulis var. higginsiiCymopterus newberryiLomatium concinnumCymopterus acaulis var. greeleyorumCymopterus acaulis var. parvusCymopterus bulbosusPteryxia terebinthina var. albiflora (3)Pteryxia terebinthina var. calcarea (3)Pteryxia petraeaPteryxia terebinthina var. calcareaCymopterus corrugatusCymopterus ibapensisCymopterus coulteriCymopterus deserticolaAletes sessiliflorusOreoxis bakeriAletes filifoliusHarbouria trachypleuraPseudocymopterus longiradiatusAletes macdouglaii subsp. macdouglaiiCymopterus beckiiPseudocymopterus montanusPteryxia davidsoniiThaspium barbinodeThaspium pinnatifidumThaspium trifoliatumZizia aureaZizia apteraPolytaenia nuttalliiPolytaenia texanaCymopterus douglassiiMusineon divaricatum var. divaricatumCymopterus macrorhizusCymopterus montanusCymopterus constancei (2)Cymopterus multinervatusCymopterus purpurascensAletes calcicolaAletes macdougalii subsp. breviradiatus (2)Aletes macdougalii subsp. breviradiatus (3)Oreoxis trotteri (2)Podistera macouniiPodistera macounii 565897818152671009464617179818770676384558210056951007873969477100

Zizia Polytaenia

Thaspium

Zizia aurea

BONAP (TAMU)Distribution Maps

ML analysis135 accessions(18 genera)

Cymopterus basalticusCymopterus gilmaniiCymopterus globosusCymopterus ripleyiLomatium juniperinumLomatium orientaleOrogenia fusiformisOrogenia linearifoliaLomatium dasycarpum subsp. dasycarpumLomatium foeniculaceum subsp. foeniculaceumCymopterus lapidosusCymopterus aboriginumCymopterus cinerariusCymopterus evertiiCymopterus panamintensis var. panamintensisOreonana purpurascensOreonana clementisLomatium cousLomatium macrocarpumOreonana vestitaLomatium grayi var. grayiLomatium grayi var. depauperatumCymopterus glaucusLomatium bradshawiiLomatium nudicauleLomatium bicolor var. bicolorLomatium triternatum subsp. platycarpumLomatium californicumLomatium idahoenseCymopterus longipesCymopterus planosusLomatium ambiguumCymopterus davisiiPteryxia terebinthina var. foeniculaceaPteryxia terebinthina var. terebinthinaPteryxia terebinthina var. californicaLomatium dissectum var. dissectumLomatium rigidumTauschia parishiiLomatium repostumLomatium shevockiiLomatium lucidumTauschia argutaCymopterus duchesnensisCymopterus goodrichiiPodistera nevadensisCymopterus minimusCymopterus purpureusCymopterus roseiCymopterus jonesiiLomatium scabrum var. scabrumLomatium parvifoliumTauschia glaucaTauschia kelloggiiLomatium howelliiCymopterus panamintensis var. acutifoliusShoshonea pulvinataTauschia texanaMusineon vaginatumOreoxis alpina subsp. alpinaLomatium greenmaniiTaenidia integerrimaLomatium latilobumLomatium graveolens var. graveolensPteryxia hendersoniiCymopterus nivalisLomatium junceumAletes humilisAletes acaulisMusineon tenuifoliumOreoxis humilisCymopterus williamsiiAletes anisatusCymopterus longilobusLomatium nuttalliiPteryxia hendersoniiLomatium brandegeiPodistera yukonensisNeoparrya lithophilaPodistera eastwoodiaeMusineon lineareCymopterus acaulis var. fendleriCymopterus acaulis var. acaulisCymopterus acaulis var. higginsiiCymopterus newberryiLomatium concinnumCymopterus acaulis var. greeleyorumCymopterus acaulis var. parvusCymopterus bulbosusPteryxia terebinthina var. albiflora (3)Pteryxia terebinthina var. calcarea (3)Pteryxia petraeaPteryxia terebinthina var. calcareaCymopterus corrugatusCymopterus ibapensisCymopterus coulteriCymopterus deserticolaAletes sessiliflorusOreoxis bakeriAletes filifoliusHarbouria trachypleuraPseudocymopterus longiradiatusAletes macdouglaii subsp. macdouglaiiCymopterus beckiiPseudocymopterus montanusPteryxia davidsoniiThaspium barbinodeThaspium pinnatifidumThaspium trifoliatumZizia aureaZizia apteraPolytaenia nuttalliiPolytaenia texanaCymopterus douglassiiMusineon divaricatum var. divaricatumCymopterus macrorhizusCymopterus montanusCymopterus constanceiCymopterus multinervatusCymopterus purpurascensAletes calcicolaAletes macdougalii subsp. breviradiatus (2)Aletes macdougalii subsp. breviradiatus (3)Oreoxis trotteri (2)Podistera macouniiPodistera macounii 565897818152671009464617179818770676384558210056951007873969477100

Cymopterus montanusCymopterus constanceiCymopterus multinervatusCymopterus purpurascens787396Cymopterus macrorhizus

Cymopterus sect. Phellopterus Nutt. ex Torr. & A. Gray

The only species group of Cymopterus that is supported, in part, by molecular evidence

Ronald HartmanBrittonia 52: 137. 2000

Cymopterus constancei

Cymopterus bulbosus

Large, showy bractlets that are basally connate and prominently veined

Showy bractlets

ML analysis135 accessions(18 genera)

Do additional data from the chloroplast genome provide increased resolution of relationships?

27 species (12 genera) examined, including:Cymopterus (6 spp.)Lomatium (8 spp.)

Angelica to root the tree

Loci Examined:• ITS• rps16 intron• trnF-trnL-trnT

Maximum Parsimony (PAUP*)

Length variationNo. of aligned positionsNo. of positions eliminatedNo. of positions constantNo. of positions informativeNo. of positions autapomorphicNo. of alignment gapsNo. of gaps parsimony informativeMaximum sequence divergence

No. of minimal length treesLength of shortest treesConsistency IndexRetention Index

ITS

437-43944473214274636.7%

>20,0001960.49550.5912

rps16

911-9769972090234411343.2%

6940.66040.8583

trnF-L-T

1693-1816188446172142752771.7%

2201450.67690.8727

Combined

3096-3229332573294411819046142.6%

164780.49260.6783

Multiple alignment of 27 DNA sequences from three loci (separately & combined)

Strict consensus trees resulting from MP analysis of ITS and rps16 intron sequence data

Angelica archangelicaZizia apteraZizia aureaThaspium trifoliatumCymopterus montanusCymopterus multinervatusCymopterus williamsiiMusineon tenuifoliumAletes anisatusCymopterus nivalisCymopterus purpureusHarbouria trachypleuraLomatium graveolensLomatium junceumMusineon divaricatumNeoparrya lithophilaPodistera eastwoodiaePteryxia hendersoniiPteryxia terebinthinaShoshonea pulvinataCymopterus globosusLomatium bicolorLomatium californicumLomatium dasycarpumLomatium juniperinumLomatium macrocarpumLomatium nudicaule967275425638 Angelica archangelicaCymopterus williamsiiHarbouria trachypleuraMusineon divaricatumMusineon tenuifoliumPteryxia hendersoniiCymopterus montanusCymopterus multinervatusZizia apteraZizia aureaThaspium trifoliatumLomatium juniperinumLomatium macrocarpumLomatium nudicauleCymopterus nivalisCymopterus purpureusLomatium dasycarpumLomatium graveolensNeoparrya lithophilaPodistera eastwoodiaePteryxia terebinthinaLomatium junceumShoshonea pulvinataLomatium bicolorLomatium californicumAletes anisatusCymopterus globosus666499868696<56022895647<599ITS rps16intron

Strict consensus trees resulting from MP analysis of trnF/L/T and combined molecular data

Angelica archangelicaCymopterus globosusLomatium graveolensCymopterus purpureusCymopterus nivalisPteryxia terebinthinaAletes anisatusLomatium dasycarpumLomatium junceumNeoparrya lithophilaPodistera eastwoodiaeShoshonea pulvinataLomatium bicolorLomatium macrocarpumLomatium juniperinumLomatium nudicauleLomatium californicumThaspium trifoliatumZizia apteraZizia aureaCymopterus montanusCymopterus multinervatusCymopterus williamsiiHarbouria trachypleuraMusineon divaricatumMusineon tenuifoliumPteryxia hendersonii695676829591689364938173 Aletes anisatusShoshonea pulvinataNeoparrya lithophilaLomatium junceumPodistera eastwoodiaeCymopterus purpureusLomatium graveolensCymopterus nivalisPteryxia terebinthinaCymopterus globosusLomatium dasycarpumLomatium bicolorLomatium californicumLomatium juniperinumLomatium macrocarpumLomatium nudicauleCymopterus williamsiiMusineon tenuifoliumPteryxia hendersoniiHarbouria trachypleuraMusineon divaricatumCymopterus montanusCymopterus multinervatusZizia apteraZizia aureaThaspium trifoliatumAngelica archangelica864440223527365568837064666866999810099trnF/L/T CombinedMolecular

Summary

• The perennial, endemic apioid umbellifers of NA (north of Mexico) constitute a weakly supported monophyletic group

• Sister to this clade are several groups of taxa, including the genus Angelica and the meso-American Arracacia clade

• Many western NA genera are not monophyletic, and the two largest genera (Cymopterus & Lomatium) are grossly polyphyletic

• Elements of Cymopterus are inextricably linked with Aletes, Lomatium, Musineon, Oreonana, Oreoxis, Orogenia, Podistera, Pseudocymopterus, Pteryxia, and Tauschia

Future Plans

• Greatest resolution of relationships was obtained by including all molecular data in a simultaneous analysis. Additional molecular data will be procured

• Cladistic analysis of morphological data will be done in an effort to uncover synapomorphies useful for generic determination. The traditional emphasis placed on fruit characters has led to highly artificial assemblages of species

• If future studies support the conclusions presented herein, and if further resolution of relationships can be achieved, radical changes to the prevailing classification will be required

Acknowledgements• NSF Grants 9407712 & 0089452

• John R. Laughnan Travel Award

• Herbert H. Ross Memorial Fund

• Francis M. and Harlie M. Clark Research Support Grants

• Program in Ecology & Evolutionary Biology Travel Grants and Summer Research Fellowship

• UIUC Graduate College Dissertation Travel Grant

• UIUC Graduate College Dissertation Completion Fellowship

• Herbarium curators for access to specimensDeborah S. Katz-Downie

Ron Hartman

Feng-Jie Sun