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Phylogenetic studies in Apiaceae tribe Oenantheae: circumscription and relationships

Phylogenetic studies in Apiaceae tribe Oenantheae: circumscription and relationships. Stephen R. Downie Krzysztof Spalik Mark F. Watson Deborah S. Katz-Downie. Outline of Presentation.

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Phylogenetic studies in Apiaceae tribe Oenantheae: circumscription and relationships

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  1. Phylogenetic studies in Apiaceae tribe Oenantheae: circumscription and relationships Stephen R. Downie Krzysztof Spalik Mark F. Watson Deborah S. Katz-Downie

  2. 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

  3. Molecular studies reveal a strongly supported Oenanthe clade

  4. Oenanthe Clade: Composition and Support 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

  5. Lilaeopsis, a simple apioid umbellifer Thick, corky ribs; no carpophore Damp, marshy or truly aquatic habitats MP analysis of rbcL, matK, & rpoC1 intron data (Petersen et al. 2002) L. mauritiana Simple, few-flowered umbels Lilaeopsis mauritiana Lilaeopsis brasiliensis (Affolter 1985) Entire, simple, linear to spatulate, septate leaves Horizontal creeping stems

  6. 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

  7. A comparison of three systems of classification • Oenantheae Dumort. (1827) • Aethusa • Coriandrum • 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 Fruits radiately ribbed • Seseleae subtribe Oenanthinae Bentham (1867) • Aethusa • Astydamia • Capnophyllum • Choritaenia • Cymbocarpum • Eurytaenia • Polemannia • Siler • Cynosciadium • Discopleura (=Ptilimnium) • Crantzia (=Lilaeopsis) • Oenanthe Ribs raised, broadly thickened at margin, corky Fruits subterete to dorsally compressed Vallecular vittae solitary Fruit anatomy (relative placement of aerenchyma) Red Highlight: Oenantheae emend. M.F. Watson & S.R. Downie

  8. Echinophoreae Scandiceae Caucalideae Coriandreae Smyrnieae (1) Hohenackerieae Pyramidoptereae Apieae (11) Angeliceae Peucedaneae (1) Tordylieae Laserpitieae Distribution of Oenantheae Genera in the Nomenclator of Pimenov & Leonov (1993)(classification modified from Drude 1898)

  9. 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

  10. Tribe Oenantheae

  11. Attributes of tribe Oenantheae Entire, simple, septate leaves Preference for wet or aquatic habitats Lilaeopsis brasiliensis Perideridia gairdneri (Oregon State University) Cicuta virosa Fascicled fibrous or tuberous roots Limnosciadium (TAMU) Lilaeopsis (Affolter 1985) Glabrous stems & leaves Cicuta (TAMU) Corky-thickened ribs (spongy cells; storage tracheids) Globose to broadly ovate fruits

  12. Oxypolis filiformis Hollow leaves with transverse septae Lilaeopsis carolinensis Oxypolis greenmanii Lilaeopsis occidentalis 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

  13. 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 • Phylogenetic Analysis: • Maximum Parsimony (PAUP*)

  14. Provisional Generic-Level Relationships in Oenantheae(strict consensus tree based on MP analysis of ITS data)

  15. Perideridia is sister to all other examined Oenantheae A possible NA origin of the tribe?

  16. Berula erecta Berula, Sium & Afrocarum are similar morphologically Sium repandum Afrocarum imbricatum Townsend, 1989, Flora of Tropical East Africa

  17. Cryptotaenia canadensis Missouri 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)

  18. Oenanthe: 40 species; Helosciadium: 5 species Oenanthe aquatica C.A.M. Lindman Flora Helosciadium nodiflorum Oenanthe nodiflora in Apium clade = Sclerosciadium 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.)

  19. Limnosciadium (BONAP) Cynosciadium (BONAP) Cynosciadium, Limnosciadium & Ptilimnium Cynosciadium digitatum Limnosciadium pinnatum Ptilimnium Ptilimnium costatum Glabrous annuals, with compound umbels Fascicled, fibrous roots Inhabits wet places in south-central and SE USA Ovoid fruits with corky thickened lateral ribs Leaves entire to compound with elongate, narrow or filiform divisions. Some species linear septate

  20. Digital Flora of Texas (TAMU) Trepocarpus aethusae Bifora americana Bifora + Neogoezia (Plunkett et al. 1996) Bifora americana Bifora americana & Trepocarpus: South-eastern USA Glabrous annuals; wetland habitats Corky fruits in Trepocarpus (that float!) Lvs pinnately decompound Trepocarpus aethusae (BONAP) Bifora americana (BONAP)

  21. Neogoezia: 5 species endemic to Mexico Constance 1987 N. macvaughii N. gracilipes Acaulescent, scapose habit Simple, many-flowered umbels Ovoid, globose fruits N. macvaughii NYBG Type Catalog Fascicled fleshy-tuberous roots Neogeozia macvaughii Usually moist habitats

  22. Perideridia: delicious tuberous roots P. gairdneri P. oregana P. howellii 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 ribs P. kelloggii

  23. Perideridia is Indigenous to USA BONAP Number of species per state: 1-3 4-5 6-7 8-9 10-11 • Strict consensus of MP trees • Perideridia neurophylla allies with Pternopetalum & Spuriopimpinella (therefore maintain as Pterygopleurum neurophyllum) P. neurophylla

  24. Lichtensteinia obscura Burtt, 1991, Edinb. J. Bot. 48(2): 225 A possible addition to tribe Oenantheae? (based on partial ITS sequence data) • Lichtensteinia • 6 spp. South Africa • 1 sp. St. Helena)

  25. Additional genera examined (or will be examined) for possible placement in tribe Oenantheae Apium Apodicarpum Asciadium Chamaele Crenosciadium Lichtensteinia Niphogeton Oreoschimperella Ottoa Pterygopleurum Pternopetalum Rutheopsis Spuriopimpinella Stoibrax Material of any of these genera would be appreciated, as well as suggestions of other taxa to survey!

  26. 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

  27. 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

  28. 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!

  29. Acknowledgements • Feng-Jie Sun • Mary Ann Feist • Changshook Lee • Carolina Calviño • Gina Colletti • NSF Grants 9407712 & 0089452 • Jean-Pierre Reduron (for material)

  30. Co-conspirators in tribe Oenantheae Mark F. Watson Krzysztof Spalik Deborah Katz-Downie

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