phylogenetic studies in apiaceae tribe oenantheae: circumscription and relationships
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Phylogenetic studies in Apiaceae tribe Oenantheae: circumscription and relationships. Stephen R. Downie Krzysztof Spalik Mark F. Watson Deborah S. Katz-Downie. Outline of Presentation. - PowerPoint PPT PresentationTRANSCRIPT
Phylogenetic studies in Apiaceae tribe Oenantheae: circumscription
and relationships
Stephen R. Downie
Krzysztof Spalik
Mark F. Watson
Deborah S. Katz-Downie
Outline of Presentation• To highlight the results of previous molecular systematic
investigations that led to the recent and modified circumscription of tribe Oenantheae
• To reveal the composition of the tribe and indicate provisional generic-level relationships based on preliminary phylogenetic analysis of rDNA ITS data
• To introduce several major genera (reduced morphology, taxonomic problems, biogeography) and discuss the implications of the phylogenetic results obtained to date
• To discuss our plans for the future
BiforaNeogoeziaSiumOenanthePerideridia90988397matK (Plunkett et al. 1996)528957100CryptotaeniaSiumBerulaCicutaOenanthePerideridiacpDNA restriction sites (Plunkett & Downie 1999)CryptotaeniaOxypolisSiumCicutaOenanthePerideridia786191rpoC1 intron(Downie et al. 1998)ITS (Downie et al. 1998, 2000)HelosciadiumBerulaSiumCryptotaeniaCicutaOenanthePerideridia<5094936010070
Molecular studies reveal a strongly supported Oenanthe clade
Region
ITS
ITS
cpDNA sites
rpl16 intron
matK
rpoC1 intron
rps16 intron
rbcL
Bootstrap (%)
100
100
100
100
97
91
86
65
Genera in Clade
Berula, Cicuta, Cryptotaenia, Oenanthe, Oxypolis, Perideridia, Sium
Berula, Cicuta, Cryptotaenia, Helosciadium, Oenanthe, Sium
Berula, Cicuta, Cryptotaenia, Oenanthe, Perideridia, Sium
Cicuta, Cryptotaenia, Oenanthe, Oxypolis Perideridia, Sium
Bifora, Neogoezia, Oenanthe, Perideridia, Sium
Cicuta, Cryptotaenia, Oenanthe, Oxypolis, Perideridia, Sium
Berula, Cicuta, Cryptotaenia, Helosciadium, Oenanthe, Sium
Cicuta, Cryptotaenia, Oenanthe, Sium
Oenanthe Clade: Composition and Support
Lilaeopsis, a simple apioid umbellifer
MP analysis of rbcL, matK, & rpoC1 intron data(Petersen et al. 2002)
Lilaeopsis brasiliensis (Affolter 1985)
Lilaeopsis mauritiana
L. mauritiana
Entire, simple, linear to spatulate, septate leavesHorizontal creeping stems
Simple, few-flowered umbels
Damp, marshy or truly aquatic habitatsThick, corky ribs; no carpophore
798694CryptotaeniaSiumNeogoeziaLilaeopsisOenanthePerideridia
The Oenanthe Clade recognized as Tribe Oenantheae Dumort.
(emend. M. F. Watson & S. R. Downie; Downie et al. 2000, 2001)
• Berula• Cicuta• Cryptotaenia• Cynosciadium• Helosciadium• Lilaeopsis• Limnosciadium
• Neogoezia• Oenanthe• Oxypolis• Perideridia• Ptilimnium• Sium
Oenantheae Dumort. (1827)• Aethusa• Coriandrum• Oenanthe
Seseleae subtribe Oenanthinae Bentham (1867)• Aethusa• Astydamia• Capnophyllum• Choritaenia• Cymbocarpum• Eurytaenia• Polemannia• Siler• Cynosciadium• Discopleura (=Ptilimnium)• Crantzia (=Lilaeopsis)• Oenanthe
Oenantheae Dumort. emend. Koso-Poljansky (1916)• Apodicarpum• Cicuta• Cymbocarpum• Cynosciadium• Cyssopetalum (=Oenanthe)• Ducrosia• Hohenackeria• Johrenia• Krubera• Oenanthe• Ptilimnium• Platylophium (=Ferulago)• Rhabdosciadium• Rhysopterus (=Cymopterus)• Stenocoelium• Thecocarpus• Trepocarpus
Red Highlight: Oenantheae emend. M.F. Watson & S.R. Downie
A comparison of three systems of classification
Fruits radiately ribbed
Ribs raised, broadly thickened at margin, corkyFruits subterete to dorsally compressedVallecular vittae solitary
Fruit anatomy(relative placement of aerenchyma)
Distribution of Oenantheae Genera in the Nomenclator of Pimenov & Leonov (1993)
(classification modified from Drude 1898)
Echinophoreae
Scandiceae
Caucalideae
Coriandreae
Smyrnieae (1)
Hohenackerieae
Pyramidoptereae
Apieae (11)
Angeliceae
Peucedaneae (1)
Tordylieae
Laserpitieae
Oenantheae Genera, Species & Distribution
Genus No. Species Distribution
Berula 2 Widespread
Cicuta 4 3 NA; 1 circumboreal
Cryptotaenia 6 Widespread
Cynosciadium 1 USA
Helosciadium 5 Eurasia
Lilaeopsis 13 New World, Australasia
Limnosciadium 2 USA
Neogoezia 5 Mexico
Oenanthe 40 Widespread
Oxypolis 7 NA
Perideridia 14 13 NA; 1 Asian
Ptilimnium 5 USA
Sium 14 Widespread
Attributes of tribe Oenantheae
Lilaeopsis (Affolter 1985)
Perideridia gairdneri(Oregon State University)
Cicuta (TAMU)
Limnosciadium (TAMU)
Globose to broadly ovate fruits
Cicuta virosa
Preference for wet or aquatic habitats
Fascicled fibrous or tuberous roots
Entire, simple, septate leaves
Corky-thickened ribs(spongy cells; storage tracheids)
Lilaeopsis brasiliensis
Glabrous stems & leaves
Oxypolis greenmanii
Oxypolis filiformis
Lilaeopsis occidentalisLilaeopsis carolinensis
Linear, septate leaves are homologous to the rachis of a normal compound leaf, with the position of septae corresponding to the position of pinnae insertion (rachis-leaves; phyllodes) The size & shape of these leaves are readily modified by the environment (Affolter 1985)
In Apiaceae, linear, septate leaves occur in: Cynosciadium, Eryngium, Lilaeopsis, Limnosciadium, Oenanthe, Ottoa, Oxypolis, Perissocoeleum & Ptilimnium
Hollow leaves with transverse septae
Materials and Methods> 260 Accessions Examined, including comprehensive representation of all 13 genera unambiguously included within tribe Oenantheae
> 60 Additional Accessions, representing taxa provisionally included within the tribe (i.e., Bifora) or those having one to several similar ecological or morphological traits (moist/wet habitat; fruits having spongy tissue; fascicled or tuberous-like roots; linear, septate or finely dissected leaves; simple umbels)
As putative outgroups, we included representation of the tribe Pleurospermeae (i.e., Aulacospermum, Physospermum, & Pleurospermum), the Komarovia clade (i.e., Hansenia, Komarovia, & Parasilaus), and Erigenia
18S nrDNA26S nrDNAITS 5ITS 3ITS 4ITS 2ITS 2ITS 15.8S215 bp224 bpITS region
Phylogenetic Analysis:• Maximum Parsimony (PAUP*)
Provisional Generic-Level Relationships in Oenantheae(strict consensus tree based on MP analysis of ITS data)
No. Species(examined / total)No. AccessionsExamined721983514CynosciadiumLimnosciadiumPtilimniumLilaeopsisBifora americanaTrepocarpusNeogeoziaAfrocarumHelosciadiumCryptotaeniaOenantheOxypolisPerideridiaBerulaSiumCicuta25823314241/11/23/55/131/11/15/51/12/214/144/43/715/409/145/55/613724= Genus is not monophyletic
Perideridia is sister to all other examined OenantheaeA possible NA origin of the tribe?
CynosciadiumLimnosciadiumPtilimniumLilaeopsisBifora americanaTrepocarpusNeogeoziaAfrocarumHelosciadiumCryptotaeniaOenantheOxypolisPerideridiaBerulaSiumCicutaUSAUSAUSANew World, AustralasiaUSAUSAMexicoAfricaEurasiaOld WorldOld WorldNANAOld WorldOld WorldNA, Circumboreal
Afrocarum imbricatum (Africa)Sium repandum (Africa)Berula thunbergii (Africa)Sium bracteatum (St.Helena)Berula erecta (Africa)Berula erecta (Europe & NA)Sium sisarumSium latifoliumSium suaveSium frigidumSium ninsiSium tenue
Berula erecta
Afrocarum imbricatumSium repandum
Townsend, 1989, Flora of Tropical East Africa
Berula, Sium & Afrocarum are similar morphologically
Cryptotaenia canadensisMissouri Flora Website
Cryptotaenia, 4-7 species
Oenantheae•N. America•Europe•Asia
Scandiceae subtribe Daucinae •Canary Islands
Pimpinella clade•Africa
“African species of Cryptotaenia are extremely close to Pimpinella” (Townsend 1989)
AfrocarumHelosciadiumCryptotaenia canadensisOenantheBerulaSiumCryptotaenia elegansMoniziaDaucusCryptotaenia africanaCryptotaenia calycinaPimpinellaCryptotaenia japonica
Oenanthe: 40 species; Helosciadium: 5 species
Oenanthe nodiflora in Apium clade
= Sclerosciadium nodiflorum
Helosciadium nodiflorumOenanthe aquatica ??Helosciadium repensHelosciadium bermejoiHelosciadium inundatumHelosciadium crassipesCryptotaeniaOenanthe (13 spp.)Oenanthe nodifloraAmmi majus
Oenanthe aquaticaC.A.M. Lindman FloraHelosciadium nodiflorum
Helosciadium was included in Apium s. lat. It comprises wetland species, with aerenchymatic tissue in its mericarps (the latter is not present in Apium s. str.)
Cynosciadium, Limnosciadium & Ptilimnium
Glabrous annuals, with compound umbelsFascicled, fibrous rootsInhabits wet places in south-central and SE USAOvoid fruits with corky thickened lateral ribsLeaves entire to compound with elongate, narrow or filiform divisions. Some species linear septateCynosciadium (BONAP)
Limnosciadium (BONAP) Ptilimnium costatumPtilimnium
Cynosciadiumdigitatum
Limnosciadiumpinnatum
CynosciadiumLimnosciadiumPtilimniumLilaeopsis
Bifora + Neogoezia(Plunkett et al. 1996)
Digital Flora of Texas (TAMU)
Bifora americana (BONAP) Trepocarpus aethusae (BONAP)
Bifora americana
Bifora americana & Trepocarpus:South-eastern USAGlabrous annuals; wetland habitatsCorky fruits in Trepocarpus (that float!)Lvs pinnately decompound
Bifora americana
Trepocarpus aethusae
Bifora americanaTrepocarpus aethusaeNeogoezia gracilipesNeogoezia breedloveiNeogoezia minorNeogoezia planipetalaNeogoezia macvaughii
Neogoezia: 5 species endemic to MexicoConstance 1987
NYBG Type Catalog
Neogeozia macvaughii
N. macvaughii
N. gracilipes
N. macvaughii
Simple, many-flowered umbels
Fascicled fleshy-tuberous roots
Acaulescent, scapose habit
Ovoid, globose fruits
Usually moist habitats
Perideridia: delicious tuberous roots P. gairdneri
P. kelloggii (Lee Dittmann)
• 14 species of glabrous perennials• Late summer to fall flowering• Unique polystelic vascular structure
(the fusion product of several
individual roots)• Moist or xerophytic habitats• Broad corky or filiform fruit ribsP. kelloggii
P. howellii
P. oregana
Perideridia is Indigenous to USA
Number of species per state:1-3 4-5 6-7 8-9 10-11
BONAP
P. neurophylla
• Strict consensus of MP trees
• Perideridia neurophylla allies with Pternopetalum & Spuriopimpinella (therefore maintain as Pterygopleurum neurophyllum)
P. bacigalupiiP. erythrorhizaP. bolanderiP. parishiiP. lemmoniiP. californicaP. pringleiP. kelloggiiP. gairdneriP. oreganaP. leptocarpaP. americanaP. howellii
A possible addition to tribe Oenantheae?(based on partial ITS sequence data)
Lichtensteinia• 6 spp. South Africa • 1 sp. St. Helena)
Burtt, 1991, Edinb. J. Bot. 48(2): 225
Lichtensteinia obscura
Additional genera examined (or will be examined) for possible placement in tribe Oenantheae
ApiumApodicarpumAsciadiumChamaeleCrenosciadiumLichtensteiniaNiphogeton
OreoschimperellaOttoaPterygopleurumPternopetalumRutheopsisSpuriopimpinellaStoibrax
Material of any of these genera would be appreciated, as well as suggestions of other taxa to survey!
Summary• Tribe Oenantheae Dumort. is a strongly supported monophyletic
group in all molecular analysis to date
• While these plants may share several morphological attributes (such as those acquired through existence in a wet environment), no unique morphological synapomorphy supports their monophyly. Some species with simplified vegetative morphology confound cladistic analysis
• Afrocarum, Trepocarpus and, possibly, Lichtensteinia, are recent additions to the tribe. The placement of Bifora americana within the tribe is confirmed
• Old World genera Berula, Cryptotaenia, Oenanthe, and Sium, as currently circumscribed, are each not monophyletic
Summary cont’d.
• The USA endemic genus Perideridia is sister taxon to all other examined Oenantheae (suggestive of a NA origin of the tribe?)
• Phytogeographical discussions based on distribution patterns of certain Oenantheae (i.e., Cryptotaenia, Perideridia & Sium) should be treated with caution prior to phylogenetic study
• Phylogenetic analysis of rDNA ITS sequences often results in fully resolved phylogenies with well-supported clades
Major Goals of Ongoing Research
• To produce a comprehensive estimate of phylogenetic relationships within tribe Oenantheae using molecular and morphological data
• To use the resultant phylogeny to elucidate patterns in the evolution of specific phenotypic characters and to explain many interesting disjunct geographic distributions
• To revise generic circumscriptions and to produce a modern classification for the group that reflects its evolutionary history. Collaborations are welcome!
Acknowledgements• Feng-Jie Sun• Mary Ann Feist• Changshook Lee• Carolina Calviño• Gina Colletti• NSF Grants 9407712
& 0089452• Jean-Pierre Reduron
(for material)