Species delimitation in recent New Zealand species radiations Heidi M. Meudt Museum of New Zealand Te Papa Tongarewa Wellington, New Zealand http://nzprn.otago.ac.nz/wiki/bin/view/NZPRN/WebHome The importance of New Zealand for understanding the phenomenon of species radiation Ilse Breitwieser, Allan Herbarium, Landcare Research, New Zealand Josephine Ward, School of Biological Sciences, University of Canterbury, New Zealand Indigenous Species Numbers • • • • • • • • • • Green algae: ~1306 Red algae: 488 Diatoms: 710 (+ ? marine) Brown algae: 153 Cyanobacteria: 336 (+ ? marine) Lichens: 1707 (388 endemic – 23%) Hepatics: 607 (314 endemic – 52%) Mosses: 483 (115 endemic – 24%) Lycophytes and ferns: 196 (90 endemic – 46%) Seed plants: 1936 (1591 endemic – 82%) Raoulia alliance Anaphalioides Leucogenes Rachelia Ewartia Raoulia Helichrysum Raoulia alliance: ecological diversity Character expressions in undescribed species “M” 1 2 3 4 5 6 7 8 9 10 Undescribed species “M” Argyrotegium Raoulia subg. Raoulia Raoulia subg. Psychrophyton Helichrysum depressum group 1. 2. 3. 4. 5. Florets mainly female Fruits papillose Distinct prostrate and uprights shoots Pappus bases not interlocking Pappus tip cells rounded 6. 7. 8. Leaves with black pigment Involucral bracts acute Pappus hairs numerous, in several series 9. Florets few 10. Leaves very small Raoulia alliance: ITS phylogeny Ewartia planchonii 85 Pterygopappus lawrencei Ewartia meredithiae 63 95 54 64 Leucogenes grandiceps Raoulia monroi Raoulia tenuicaulis Raoulia sp. ‘M’ Helichrysum intermedium Leucogenes leontopodium Helichrysum filicaule Raoulia rubra Helichrysum lanceolatum Anaphalioides mariae Anaphalioides trinervis Anaphalioides subrigida Anaphalioides hookeri Anaphalioides alpina Anaphalioides bellidioides Ozothamnus leptophyllus Anaphalis triplinervis 92 63 79 100 63 Breitwieser et al., 1999 Ewartia sinclairii Rachelia glaria “NZ clade”: Anaphalioides Ewartia* Helichrysum* Leucogenes Rachelia Raoulia* Anaphalis javanica Pseudognaphalium luteoalbum outgroups Craspedia Small sequence variation within the NZ lineage Neighbour network from AFLP profiles of a wide sample of NZ Craspedia 83 New Zealand NZ 100 Craspedia 96 AUS 3 100 100 99 94 100 AUS 2 99 100 AUS 1 100 Helichrysum lanceolatum Pynosorus globosus Pynosorus chrysanthes Stuartina muelleri 5 changes ITS & ETS sequences (nuclear) Brachyglottis Brachyglottis repanda Brachyglottis saxifragoides Brachyglottis cassinioides Brachyglottis cassinioides Brachyglottis alliance ITS 98 58 85 100 91 96 92 100 100 94 53 65 51 69 100 43 92 59 95 85 46 81 29 Brachyglottis bellidioides Brachyglottis haastii Brachyglottis southlandica Brachyglottis compacta Brachyglottis greyi Bedfordia linearis Bedfordia salicina Brachyglottis brunonis Brachyglottis cassinioides Brachyglottis huntii Haastia recurva Haastia sinclairii Brachyglottis adamsii Brachyglottis arborescens Brachyglottis elaeagnifolia Brachyglottis kirkii Brachyglottis perdicioides Brachyglottis repanda Brachyglottis sciadophila Dolichoglottis scorzoneroides Haastia pulvinaris Traversia baccharoides Senecio dunedinensis Senecio quadridentatus Senecio glaucophyllus Senecio rufiglandulosus Senecio lautus Senecio vulgaris Senecio glomeratus Senecio minimus Senecio jacobaea Petasites fragrans Euchiton fordianus Inula orientalis Pachystegia insignis Abrotanella muscosa Cotula coronopifolia Calendula officinalis Dasyphyllum dicanthoides 5' trnK / matK Brachyglottis bellidioides Brachyglottis haastii Brachyglottis southlandica Brachyglottis compacta Brachyglottis greyi Bedfordia linearis Bedfordia salicina Brachyglottis brunonis Brachyglottis cassinioides Brachyglottis huntii Haastia recurva Haastia sinclairii Brachyglottis adamsii Brachyglottis arborescens Brachyglottis elaeagnifolia Brachyglottis kirkii Brachyglottis perdicioides Brachyglottis repanda Brachyglottis sciadophila Dolichoglottis scorzoneroides Haastia pulvinaris Traversia baccharoides Senecio dunedinensis Senecio quadridentatus Senecio glaucophyllus Senecio rufiglandulosus Senecio minimus Senecio vulgaris Senecio glomeratus Senecio lautus Senecio jabcobaea Petasites fragrans Euchiton fordianus Inula orientalis Pachystegia insignis Abrotanella muscosa Cotula coronopifolia Calendula officinalis Dasyphyllum dicanthoides 62 88 73 100 63 62 86 55 Brachyglottis alliance: Brachyglottis, Dolichoglottis, Traversia, Haastia, Bedfordia Traversia baccharoides Haastia pulvinaris Brachyglottis cassinioides Haastia sinclairii Dolichoglottis lyallii Myosotis Winkworth et al. 2002 Mol.Phyl.Evol. Myosotis ITS, matK, psbA-trnA, ndhF New Zeland Eurasia Ourisia Ourisia Meudt & Simpson. 2006. Biol. J. Linn. Soc. Veronica Veronica evenosa Veronica tetragona Veronica stricta Veronica pulvinaris Veronica tubata Veronica lilliputiana Veronica hulkeana Veronica elliptica Veronica cupressoides Veronica hookeriana Veronica tetragona Veronica salicifolia Veronica chionohebe Veronica raoulii Veronica venustula Veronica speciosa Slide courtesy of Phil Garnock-Jones Albach & Meudt (2010); Evonne Low, thesis hebe clade sun hebe clade Veronica speedwell hebe clade snow hebe clade semiwhipcord hebe clade Ranunculus R. sericophyllus R. pachyrrhizus Carlos Lehnebach 2008: Phylogenetic affinities, species delimitation and adaptive radiation of New Zealand Ranunculus. (PhD thesis) Species radiation Evolution from single found population into array of morphologically and ecologically diverse species McGlone, M., Stevens, G. & McCulloch, B. Prehistoric New Zealand. 1988 Heinemann Reed, Auckland. New Zealand Plant Radiation Network Species radiation: • Significance of polyploidy • Species delimitation • Impact of hybridisation • Adaptive radiations • Drivers of plant diversification Polyploidy • Polyploidy common in New Zealand flora, e.g., Raoulia alliance, Veronica • Polyploidy common in New Zealand plant radiations Three ploidy levels: Leucogenes L. grandiceps, L. leontopodium: diploid L. neglecta: tetraploid L. tarahaoa: octoploid L. leontopodium L. grandiceps L. neglecta insert map here L. tarahaoa Principal components analysis of AFLP data for Leucogenes •AFLP profiles group samples of each species. L. grandiceps L. neglecta •Diploid L. leontopodium & tetraploid L. neglecta cluster together. This is consistent with derivation of L. neglecta from L. leontopodium by autopolyploidy. •Octoploid L. tarahoa does not cluster with either L. neglecta or L. leontopodium. This indicates that the species has independent origin. L. leontopodium L. tarahaoa Pachycladon Myosotis New Zealand Ozothamnus Raoulia : diversity of mat and cushion form Hybrids Raoulia mammilaris X Leucogenes grandiceps Adaptive radiation: Ranunculus lyallii M. pulvinaris Adaptive radiation: Myosotis M. traversii M. colensoi Ecology of plant radiations in New Zealand Are our plant radiations adaptive and are there adaptive zones? Within radiations is the pattern generalist ancestors, specialised descendants? Do radiations interact so that colonisation chronology impacts on niche dominance and habitat use? Conclusion New Zealand ideal for study of plant evolution • Oceanic island • Isolation • Continental fragment • Long fossil record • Diverse flora with Gondwana elements and origin from long distance dispersal • Dispersal followed by radiation and range expansion Acknowledgements Allan Herbarium staff, Landcare Research NZPRN members Photos: Phil Garnock-Jones, Peter Heenan, Carlos Lehnebach, Pete Lockhart, Heidi Meudt, John Lovis, and Rainer Vogt http://nzprn.otago.ac.nz/wiki/bin/view/NZPRN/WebHome