Evolutionary significance of long-distance dispersal and hybridisation in the New Zealand endemic genus Hoheria (Malvaceae)
Steven J. Wagstaff A D , Brian P. J. Molloy B and Jennifer A. Tate CA Allan Herbarium, Landcare Research, PO Box 40, Lincoln 7640, New Zealand.
B Research Associate, Landcare Research, PO Box 40, Lincoln 7640, New Zealand.
C Massey University, Institute of Molecular Biosciences, Private Bag 11222, Palmerston North, New Zealand.
D Corresponding author. Email: wagstaffs@landcareresearch.co.nz
Australian Systematic Botany 23(2) 112-130 https://doi.org/10.1071/SB09017
Submitted: 22 April 2009 Accepted: 25 January 2010 Published: 31 May 2010
Abstract
Parsimony and SplitsTree analyses of ITS and 5′ trnK/matK sequence data were used to assess the extent of hybridisation and its evolutionary significance in the New Zealand endemic genus Hoheria A.Cunn. The seven species of Hoheria form a monophyletic group along with the New Zealand endemic genus Plagianthus J.R.Forst. & G.Forst. and the Australian endemics Asterotrichion Klotzsch. and Gynatrix Alef. This strongly supported clade is sister to a clade of Australian species of Lawrencia Hook. The inferred evolutionary history of Hoheria suggests that the extant species are derived from a common ancestor that arrived in New Zealand by long-distance dispersal. There was little divergence among the species of Hoheria in either their ITS or trnK/matK sequences. Two known Hoheria hybrids exhibited overlapping heteromorphic nucleotides at virtually all of the variable positions. Approximately 40% of the other Hoheria accessions in our study retain similar heteromorphic sites. These polymorphisms were shared among the deepest branches in the ITS phylogeny, which potentially suggests that hybridisation has occurred throughout the evolutionary history of Hoheria. The phylogenetic structure of the ITS phylogeny completely collapsed in the strict consensus tree, and there was significant conflict between the biparentally inherited ITS phylogeny and the maternally inherited trnK/matK phylogeny. However, the removal of known and suspected hybrids resulted in parsimony trees that were more resolved. SplitsTree analyses revealed incompatible signals in the data, but recovered well supported groups that diverged from a central boxy network. Although the species of Hoheria are isolated by their ecological preferences or geographical distributions, interspecific hybrids are common in urban areas where the species are often planted.
Acknowledgements
The authors gratefully acknowledge the assistance of A. Markey in collecting Hoheria specimens; W. G. Lee, D. Glenny, B. Clarkson, P. de Lange, M. Morgan-Richards, J. Young, and anonymous reviewers for their insightful comments and advice; and D. Lee for valuable information about the fossil history of Hoheria and Plagianthus in New Zealand. We also thank three anonymous reviewers for their helpful manuscript suggestions. This research was supported by the New Zealand Foundation for Research, Science and Technology through the Defining New Zealand’s Land Biota OBI.
Allan HH
(1926a) Illustrations of wild hybrids in the New Zealand flora II. Genetica 8, 369–374.
| Crossref | GoogleScholarGoogle Scholar |
Allan HH
(1926b) Epharmonic response in certain New Zealand species and its bearing on taxonomic questions. Journal of Ecology 14, 72–91.
| Crossref | GoogleScholarGoogle Scholar |
Alvarez I, Wendel JF
(2003) Ribosomal ITS sequences and plant phylogenetic inference. Molecular Phylogenetics and Evolution 29, 417–434.
| Crossref | GoogleScholarGoogle Scholar |
CAS |
PubMed |
Baldwin BG,
Sanderson MJ,
Porter JM,
Wojciechowski MF,
Campbell CS, Donoghue MJ
(1995) The ITS region of nuclear ribosomal DNA: a valuable source of evidence on angiosperm phylogeny. Annals of the Missouri Botanical Garden 82, 247–277.
| Crossref | GoogleScholarGoogle Scholar |
Ballard HE, Sytsma KJ
(2000) Evolution and biogeography of the woody Hawaiian violets (Viola, Violaceae): arctic origins, herbaceous ancestry, and bird dispersal. Evolution 54, 1521–1532.
| Crossref | GoogleScholarGoogle Scholar | PubMed |
Barreda V
(1993) Late Oligocene?–Miocene pollen of the families Compositae, Malvaceae, and Polygonaceae from the Chenque formation, Golfo San Jorge Basin, south-eastern Argentina. Palynology 17, 169–186.
Barreda V,
Anzótegui LM,
Prieto AR,
Aceñolaza P, Bianchi MM , et al.
(2007) Diversificación y cambios de las angiosperms durante el Neógeno en Argentina. Asociación Paleontológica Argentina 11, 173–191.
Bayer RJ, Starr JR
(1998) Tribal phylogeny of the Asteraceae based on two non-coding chloroplast sequences, the trnL intron and trnL–trnF intergenic spacer. Annals of the Missouri Botanical Garden 85, 242–256.
| Crossref | GoogleScholarGoogle Scholar |
Böhle UR,
Hilger HH, Martin WF
(1996) Island colonization and evolution of the insular woody habit in Echium L. (Boraginaceae). Proceedings of the National Academy of Sciences, USA 93, 11 740–11 745.
| Crossref | GoogleScholarGoogle Scholar |
Carlquist S
(1966) The biota of long-distance dispersal. IV. Genetic systems in the floras of oceanic islands. Evolution 20, 433–455.
| Crossref | GoogleScholarGoogle Scholar |
Cockayne L
(1901) An inquiry into the seedling forms of New Zealand phanerogams and their development. Transactions and Proceedings of the New Zealand Institute
, 263–298.
Cockayne L
(1906) Notes on the subalpine scrub of Mount Fyffe (Seaward Kaikouras). Transactions and Proceedings of the New Zealand Institute 38, 361–374.
Cockayne L, Allan HH
(1926) Notes on New Zealand floristic botany, including descriptions of new species. Transactions and Proceedings of the New Zealand Institute 56, 21–33.
Corriveau JL, Coleman AW
(1988) Rapid screening method to detect potential biparental inheritance of plastid DNA and results for over 200 angiosperm species. American Journal of Botany 75, 1443–1458.
| Crossref | GoogleScholarGoogle Scholar |
Cowie RH, Holland BS
(2006) Dispersal is fundamental to biogeography and the evolution of biodiversity on oceanic islands. Journal of Biogeography 33, 193–198.
| Crossref | GoogleScholarGoogle Scholar |
Cronk QC,
Kiehn M,
Wagner WL, Smith JF
(2005) Evolution of Cyrtandra (Gesneriaceae) in the Pacific Ocean: the origin of a supertramp. American Journal of Botany 92, 1017–1024.
| Crossref | GoogleScholarGoogle Scholar |
Dawson MI, Beuzenberg EJ
(2000) Index of chromosome numbers of indigenous New Zealand spermatophytes. New Zealand Journal of Botany 38, 47–150.
de Lange PJ, Cameron EK
(1999) The vascular flora of Aorangi Island, Poor Knights Islands, Northern New Zealand. New Zealand Journal of Botany 37, 433–468.
de Lange PJ,
Norton DA,
Heenan PB,
Courtney SP,
Molloy BPJ,
Ogle CC, Rance BD
(2004) Threatened and uncommon plants of New Zealand. New Zealand Journal of Botany 42, 45–76.
Degnan JH, Rosenberg NA
(2009) Gene tree discordance, phylogenetic inference and the multispecies coalescent. Trends in Ecology & Evolution 24, 332–340.
| Crossref | GoogleScholarGoogle Scholar |
Dettmann ME, Clifford HT
(2000) Monocotyledon fruits and seeds, and an associated palynoflora from Eocene–Oligocene sediments of coastal central Queensland, Australia. Review of Palaeobotany and Palynology 110, 141–173.
| Crossref | GoogleScholarGoogle Scholar | PubMed |
Doyle JJ, Doyle JL
(1987) A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochemical Bulletin 19, 11–15.
Dunbar-Co S,
Wieczorek AM, Morden CW
(2008) Molecular phylogeny and adaptive radiation of the endemic Hawaiian Plantago species (Plantaginaceae). American Journal of Botany 95, 1177–1188.
| Crossref | GoogleScholarGoogle Scholar |
CAS |
Felsenstein J
(1985) Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39, 783–791.
| Crossref | GoogleScholarGoogle Scholar |
Francisco-Ortega J,
Jansen RK, Santos-Guerra A
(1996) Chloroplast DNA evidence of colonization, adaptive radiation, and hybridization in the evolution of the Macronesian flora. Proceedings of the National Academy of Sciences, USA 93, 4085–4090.
| Crossref | GoogleScholarGoogle Scholar |
CAS |
Groves BE, Hair JB
(1971) Contributions to a chromosome atlas of the New Zealand Flora – 15 miscellaneous familes. New Zealand Journal of Botany 9, 569–579.
Heads M
(2000) A new species of Hoheria (Malvaceae) from the Poor Knights Islands and Hen and Chickens Islands, New Zealand. New Zealand Journal of Botany 38, 373–377.
Heads M
(2009) Inferring biogeographic history from molecular phylogenies. Biological Journal of the Linnean Society. Linnean Society of London 98, 757–774.
| Crossref | GoogleScholarGoogle Scholar |
Heenan PB,
Dawson MI,
Redmond DN, Wagstaff SJ
(2005) Relationships of the New Zealand mountain ribbonwoods (Hoheria glabrata and H. lyallii: Malvaceae), based on molecular and morphological data. New Zealand Journal of Botany 43, 527–549.
Hillis D,
Moritz C,
Porter CA, Baker RJ
(1991) Evidence for biased gene conversion in concerted evolution of ribosomal DNA. Science 251, 308–310.
| Crossref | GoogleScholarGoogle Scholar |
CAS |
PubMed |
Howarth D, Baum DA
(2005) Genealogical evidence of homoploid hybrid speciation in an adaptive radiation of Scaevola (Goodeniaceae) in the Hawaiian Islands. Evolution 59, 948–961.
|
CAS |
PubMed |
Huson DH
(1998) SplitsTree: analyzing and visualizing evolutionary data. Bioinformatics 14, 68–73.
| Crossref | GoogleScholarGoogle Scholar |
CAS |
PubMed |
Huson DH, Bryant D
(2006) Application of phylogenetic networks in evolutionary studies. Molecular Biology and Evolution 23, 254–267.
| Crossref | GoogleScholarGoogle Scholar |
CAS |
PubMed |
Iglesias AP,
Wilf P,
Johnson MA,
Zamuner AB,
Cúneo NR, Matheos SD
(2007) A Paleocene lowland macroflora from Patagonia reveals significantly greater richness than North American analogs. Geology 35, 947–950.
| Crossref |
Joly S,
McLenachan PA, Lockhart PJ
(2009) A statistical approach for distinguishing hybridization and incomplete lineage sorting. American Naturalist 174, E54–E70.
| Crossref | GoogleScholarGoogle Scholar | PubMed |
Jorgensen T, Olesen JM
(2001) Adaptive radiation of island plants: evidence from Aeonium (Crassulaceae) of the Canary Islands. Perspectives in Plant Ecology and Evolution 4, 29–42.
| Crossref | GoogleScholarGoogle Scholar |
Kubatko LS
(2009) Identifying hybridization events in the presence of coalescence via model selection. Systematic Biology 58, 478–488.
| Crossref | GoogleScholarGoogle Scholar |
CAS |
Lander NS
(1984) Revision of the Australian genus Lawrencia Hook. (Malvaceae: Malveae). Nuytsia 5, 201–271.
Landis CA,
Campbell HJ,
Begg JG,
Mildenhall DC,
Patterson AM, Trewick SA
(2008) The Waipounamu erosion surface: questioning the antiquity of the New Zealand land surface and terrestrial flora and fauna. Geological Magazine 145, 173–197.
| Crossref | GoogleScholarGoogle Scholar |
Linder CR, Rieseberg LH
(2004) Reconstructing patterns of reticulate evolution in plants. American Journal of Botany 91, 1700–1708.
| Crossref | GoogleScholarGoogle Scholar | PubMed |
Lindqvist C,
Motley TJ,
Jeffrey JJ, Albert VA
(2003) Cladogenesis and reticulation in the Hawaiian endemic mints. Cladistics 19, 480–495.
| Crossref | GoogleScholarGoogle Scholar |
Macphail MK
(1997) Comment on M. Pole (1994): The New Zealand flora – entirely long-distance dispersal. Journal of Biogeography 22, 625–635.
Manchester SR
(1992) Flowers, fruit, and pollen of Florissantia, an extinct malvalean genus from the Eocene and Oligocene of Western North America. American Journal of Botany 79, 996–1008.
| Crossref | GoogleScholarGoogle Scholar |
Manchester SR
(1994) Inflorescence bracts of fossil and extant Tilia in North America, Europe, and Asia: patterns of morphologic divergence and biogeographic history. American Journal of Botany 81, 1176–1185.
| Crossref | GoogleScholarGoogle Scholar |
Mautino LR,
Cuadrado GA, Anzótegui LM
(2004) Novedades taxonómicas, diversida y significado evolutivo del pollen de Malvaceae en el Terciario de Argentina. Revista Española de Micropaleotologia 36, 467–483.
McBreen K, Lockhart PJ
(2006) Reconstructing reticulate evolutionary histories in plants. Trends in Plant Science 11, 398–404.
| Crossref | GoogleScholarGoogle Scholar |
CAS |
PubMed |
McDade LA
(1992) Hybrids and phylogenetic systematics II. The impact of hybrids on cladistic analysis. Evolution 46, 1329–1346.
| Crossref | GoogleScholarGoogle Scholar |
McGlone MS
(1983) The Polynesian deforestation of New Zealand: a preliminary synthesis. Archaeology in Oceania 18, 11–25.
McGlone MS,
Dungan RJ,
Hall GM, Allen RB
(2004) Winter leaf loss in the New Zealand woody flora. New Zealand Journal of Botany 42, 1–19.
McLoughlin S
(2001) The breakup history of Gondwana and its impact on pre-Cenozoic floristic provincialism. Australian Journal of Botany 49, 271–300.
| Crossref | GoogleScholarGoogle Scholar |
McQueen DR
(1954) Fossil leaves, fruits and seeds from the Wanganui Series (Plio-Pleistocene) of New Zealand. New Zealand Journal of Botany 82, 667–676.
Melville R
(1966) Contributions to the Flora of Australia: VII. Generic delimitation in the Plagianthus complex. Kew Bulletin 20, 511–516.
| Crossref | GoogleScholarGoogle Scholar |
Meng C, Kubatko LS
(2009) Detecting hybrid speciation in the presence of incomplete lineage sorting using gene tree incongruence: a model. Theoretical Population Biology 75, 35–45.
| Crossref | GoogleScholarGoogle Scholar | PubMed |
Mildenhall DC
(1980) New Zealand late Cretaceous and Cenozoic plant biogeography: a contribution. Palaeogeography, Palaeoclimatology, Palaeoecology 31, 197–233.
| Crossref | GoogleScholarGoogle Scholar |
Molloy BPJ,
Burrows CJ,
Cox JE,
Johnston JA, Wardle P
(1963) Distribution of subfossil forest remains, eastern South Island, New Zealand. New Zealand Journal of Botany 1, 68–77.
Morgan-Richards M,
Smissen RD,
Shepherd LD,
Wallis GP,
Hayward JJ,
Chan C-H,
Chambers GK, Chapman HM
(2009) A review of genetic analyses of hybridisation in New Zealand. Journal of the Royal Society of New Zealand 39, 15–34.
Muller J
(1984) Significance of fossil pollen for angiosperm history. Annals of the Missouri Botanical Garden 71, 419–443.
| Crossref | GoogleScholarGoogle Scholar |
Nathan R,
Schurr FM,
Spiegel O,
Steinitz O,
Trakhtenbrot A, Tsoar A
(2008) Mechanisms of long-distance seed dispersal. Trends in Ecology & Evolution 23, 638–647.
| Crossref | GoogleScholarGoogle Scholar |
Newstrom L, Robertson A
(2005) Progress in understanding pollination systems in New Zealand. New Zealand Journal of Botany 43, 1–59.
Oliver WBR
(1925) Vegetation of the Poor Knights Islands. New Zealand Journal of Science and Technology 7, 371–384.
Pocknall DT
(1982) Palynology of late Oligocene Pomahaka Estuarine Bed sediments, Waikoikoi, Southland, New Zealand. New Zealand Journal of Botany 20, 263–287.
Pole M
(1994) The New Zealand flora – entirely long-distance dispersal? Journal of Biogeography 21, 625–635.
| Crossref | GoogleScholarGoogle Scholar |
Posada D, Crandall KA
(2001) Intraspecific gene genealogies: trees grafting into networks. Trends in Ecology & Evolution 16, 37–45.
| Crossref | GoogleScholarGoogle Scholar |
Sakai AK,
Weller WL,
Wagner WL,
Nepokroeff M, Culley TM
(2006) Adaptive radiation and evolution of breeding systems in Schiedea (Caryophyllaceae), an endemic Hawaiian genus. Annals of the Missouri Botanical Garden 93, 49–63.
| Crossref | GoogleScholarGoogle Scholar |
Sang T, Zhong Y
(2000) Testing hybridization hypotheses based on incongruent gene trees. Systematic Biology 49, 422–434.
| Crossref | GoogleScholarGoogle Scholar |
CAS |
PubMed |
Seehausen O
(2004) Hybridization and adaptive radiation. Trends in Ecology & Evolution 19, 198–207.
| Crossref | GoogleScholarGoogle Scholar |
Simmons MP, Ochoterena H
(2000) Gaps as characters in sequence-based phylogenetic analyses. Systematic Biology 49, 369–381.
| Crossref | GoogleScholarGoogle Scholar |
CAS |
PubMed |
Soltis DE,
Mavrodiev EV,
Doyle JJ,
Rauscher J, Soltis PS
(2008) ITS and ETS sequence data and phylogeny reconstruction in allopolyploids and hybrids. Systematic Botany 33, 7–20.
| Crossref | GoogleScholarGoogle Scholar |
Sprague TA, Summerhayes VS
(1926) The taxonomic position of Hoheria lyallii. Kew Bulletin of Miscellaneous Information 1926, 214–220.
| Crossref | GoogleScholarGoogle Scholar |
Tate JA,
Fuertes Aguilar J,
Wagstaff SJ,
La Duke JC,
Slotta TAB, Simpson BB
(2005) Phylogenetic relationships within the tribe Malveae (subfamily Malvoideae, Malvaceae) as inferred from ITS sequence data. American Journal of Botany 92, 584–602.
| Crossref | GoogleScholarGoogle Scholar |
CAS |
Templeton AR
(2008) The reality and importance of founder speciation in evolution. BioEssays 30, 470–479.
| Crossref | GoogleScholarGoogle Scholar | PubMed |
Thompson JD,
Gibson TJ,
Plewniak F,
Jeanmougin F, Higgins DG
(1997) The Clustal X windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research 25, 4876–4882.
| Crossref | GoogleScholarGoogle Scholar |
CAS |
PubMed |
Wagstaff SJ, Breitwieser I
(2004) Phylogeny and classification of Brachyglottis (Senecioneae, Asteraceae): an example of a rapid species radiation in New Zealand. Systematic Botany 29, 1003–1010.
| Crossref | GoogleScholarGoogle Scholar |
Wardle P
(1985) Environmental influences on the vegetation of New Zealand. New Zealand Journal of Botany 23, 773–788.
Wilf P,
Johnson KR,
Cúneo NR,
Smith ME,
Singer BS, Gandolfo MA
(2005) Eocene plant diversity at Laguna del Hunco and Río Pichileufú, Patagonia, Argentina. American Naturalist 165, 634–650.
| Crossref | GoogleScholarGoogle Scholar | PubMed |
Winkworth RC,
Robertson AW,
Ehrendorfer F, Lockhart PJ
(1999) The importance of dispersal and recent speciation in the flora of New Zealand. Journal of Biogeography 26, 1323–1325.
| Crossref | GoogleScholarGoogle Scholar |
Winkworth RC,
Wagstaff SJ,
Glenny D, Lockhart PJ
(2002) Plant dispersal N.E.W.S. from New Zealand. Trends in Ecology & Evolution 17, 514–520.
| Crossref | GoogleScholarGoogle Scholar |
Winkworth RC,
Wagstaff SJ,
Glenny D, Lockhart PJ
(2005) Evolution of the New Zealand mountain flora: Origins, diversification and dispersal. Organisms, Diversity & Evolution 5, 237–247.
| Crossref | GoogleScholarGoogle Scholar |
Wissemann V
(2007) Plant evolution by means of hybridization. Systematics and Biodiversity 5, 243–253.
| Crossref | GoogleScholarGoogle Scholar |
Zamaloa MC, Romero RJ
(1990) Some spores and pollen from the Cullen Formation (Upper Eocene to Middle Oligocene), Tierra del Fuego, Argentina. Palynology 14, 123–133.
Appendix 1. Species, collection locality, DNA accession, herbarium accession, GenBank accession: ITS, 5′trnK/matK
Asterotrichion discolor (Hook.f.) Melville, Australia, Tasmania, Bluff River, A. Moscal 8224, 1.vi.1984, 9.72, CHR 491243, AY591811, GU045813. Gynatrix pulchella (Willd.) Alef., Australia, Canberra, CSIRO grounds, lower eastern slopes of Black Mountain, R. Pullen 8529, 14.x.1973, 9.69, CHR 410595A, AY591826, GU045814. Hoheria × allanii (Cockayne) Heenan, Cultivated, ex New Zealand, Canterbury, near Ashburton, S. J. Wagstaff, i.2006, 6.02, CHR 580149, GU045775, GU045797. Hoheria angustifolia Raoul, New Zealand, Otago, Otago Peninsula, Taiaroa Bush, A. Markey, 11.i.2000, 0.10, CHR 534905A, AY944585, AY944608. Hoheria angustifolia Raoul, New Zealand, Canterbury, Peel Forest, M. I. Dawson and S. J. Wagstaff, 13.iii.2002, 2.65, CHR 559132, GU045771, GU045793. Hoheria angustifolia Raoul, New Zealand, Wellington, Wainuioru Valley, Wainuioru, ‘Ngatapa Bush’, P. J. de Lange 2189 and T. Silbery, 29.viii.1993, 9.29, CHR 497780A, GU045773, GU045793. Hoheria angustifolia Raoul, New Zealand, Southland, Manse Bush, G. M. Rogers, 18.xii.1993, 9.64, CHR 500064, GU045772, GU045795. Hoheria angustifolia Raoul, New Zealand, Canterbury, Banks Peninsula, Lake Forsyth, D. R. Given 12172, 15.i.1980, 9.65, CHR 495145, GU045774, GU045794. Hoheria angustifolia Raoul × Hoheria sexstylosa Colenso, New Zealand, Canterbury, Port Hills, Marleys Hill, P. B. Heenan, B. P. J. Molloy and S. J. Wagstaff, 6.ix.2005, 5.59, CHR 95703, GU045776, GU045798. Hoheria equitum Heads, Cultivated, ex New Zealand, North Auckland, Poor Knights Islands, Tawhiti Rahi, P. J. de Lange 6543, 23.vii.2005, 5.47, AK 291559, GU045777, GU045799. Hoheria equitum Heads, New Zealand, Hen Island, D. Town, 11.i.2006, 6.01, CHR 503902, GU045778, GU045800. Hoheria equitum Heads, Cultivated, ex New Zealand, North Auckland, Poor Knights Islands, Aorangi Island, P. B. Heenan, 17.ix.1999, 9.54, CHR 529193, AY944586, AY944609. Hoheria glabrata Sprague & Summerh., New Zealand, Canterbury, Arthur’s Pass NP, Otira Valley, S. J. Wagstaff, 9.iii.2001, 1.18, CHR 541761, AY944588, AY944611. Hoheria glabrata Sprague & Summerh., New Zealand, Taranaki, A. P. Druce, Mar. 1964, 2.145, CHR 159482, AY944592, AY944615. Hoheria glabrata Sprague & Summerh., New Zealand, Fiordland, upper Hollyford Valley, P. N. Johnson 699, 19.i.1988, 9.30, CHR 439552, AY944589, AY944612. Hoheria glabrata Sprague & Summerh., New Zealand, Otago, Pisa Range, Lochar Burn, P. Wardle 94/184, 24.iv.1994, 9.66, CHR 508679B, AY944590, AY944613. Hoheria lyallii Hook.f., New Zealand, Canterbury, Seaward Kaikoura Range, Kaikoura, Mt Fyffe Hut, A. Markey, 17.iv.2002, 2.11, CHR 560121, AY944600, AY944623. Hoheria lyallii Hook.f., New Zealand, Nelson, Turk’s Cap Range, A. P. Druce, ii.1989, 2.13, CHR 395652, AY944602, AY944625. Hoheria lyallii Hook.f., New Zealand, Canterbury, Peel Forest, Raules Gully Scenic Reserve, S. J. Wagstaff and M. I. Dawson, 13.iii.2002, 2.68, CHR 559144, AY944599, AY944622. Hoheria lyallii Hook.f., New Zealand, Canterbury, Boundary Stream, main road to Hermitage, S. J. Wagstaff and M. I. Dawson, 12.iii.2002, 2.80, CHR 559129, AY944594, AY944617. Hoheria ovata G.Simpson & J.S.Thomson, New Zealand, Nelson, Mt Burnett, P. B. Heenan and P. J. de Lange, 8.iv.1997, 2.147, CHR 512522A, AY944603, AY944626. Hoheria ovata G.Simpson & J.S.Thomson, New Zealand, Westland, Fraser Ck, F. B. Overmars, 24.viii.2000, 2.149, CHR 536190, GU045780, GU045802. Hoheria ovata G.Simpson & J.S.Thomson, New Zealand, Westland, Punakaiki, Dolomite Point, P. J. de Lange 6547, 24.vii.2005, 5.51, AK 291563, GU045779, GU045801. Hoheria populnea A.Cunn., Cultivated, ex New Zealand, North Auckland, Bream Head, P. B. Heenan, ii.2002, 2.54, CHR 560114, GU045782, GU045804. Hoheria populnea A.Cunn., New Zealand, Great Barrier Island, P. B. Heenan and P. J. de Lange, 6.x.2005, 5.79, CHR 574260, GU045781, GU045803. Hoheria populnea A.Cunn. Cultivated, New Zealand, Canterbury, Lincoln, unknown origin, S. J. Wagstaff, 15.ii.1999, 9.27, CHR 529983, AY944604, AY944627. Hoheria sexstylosa Colenso, New Zealand, Canterbury, Banks Peninsula, above Governors Bay, H. D. Wilson BP406, 31.vii.1984, 9.51, CHR 479448, GU045784, GU045806. Hoheria sexstylosa Colenso, New Zealand, Hawke’s Bay, Waiwhare, Taihape Rd, B. Macmillan, 15.xii.1982, 1.38, CHR 446836, AY944605, AY944628. Hoheria sexstylosa Colenso, New Zealand, Canterbury, Banks Peninsula, Kennedys Bush, P. B. Heenan, B. P. J. Molloy and S. J. Wagstaff, 6.ix.2005, 5.63, CHR 595723, GU045783, GU045805. Hoheria sexstylosa Colenso, cultivated, ex New Zealand, Canterbury, Banks Peninsula, Kaituna Reserve, S. J. Wagstaff, 15.ii.1999, 9.25, CHR 529984, GU045785, GU045807. Hoheria ‘Tararua’, New Zealand, Wellington, Tararua Range, Mt Bruce, 8.50, AK 304592, GU045789, GU045808. Hoheria ‘Tararua’, New Zealand, Wellington, Tararua Range, Mt Bruce, 8.51, AK 304593, GU045788, GU045810. Hoheria ‘Tararua’, New Zealand, Wellington, Tararua Range, Mt Bruce, 8.53, AK 304594, GU045790, GU045811. Hoheria ‘Tararua’, New Zealand, Wellington, Tararua Range, P. B. Heenan and P. J. de Lange, 1.v.2003, 3.3, CHR 565512, GU045787, GU045809. Hoheria ‘Tararua’, New Zealand, Wellington, Hikawera, P. J. de Lange 2202 and T. Silbery, 29.viii.1993, 9.52, CHR 497795A, GU045786, GU045812. Lawrencia glomerata Hook., Australia, South Australia, Lake Torrens Basin, Carrapateena Arm, Salt Ck, south of Archie Beavis Dam, J. Z. Weber 1321, 5.ix.1968, 9.70, CHR 380446A, AY591836, GU045815. Lawrencia helmsii (F.Muell. & Tate) Lander, Australia, Western Australia, Lake Austin, L. A. Craven 5033, 22.iv.1978, 9.68, CHR 380678, AY591852, GU045816. Lawrencia spicata Hook., Australia, Tasmania, Dorans Rd, near Lauderdale, A. M. Buchanan 4540, 28.xi.1984, 9.71, CHR 423878, AY591835, GU045817. Lawrencia squamata Miq., Australia, South Australia, Nullarbor Region, N. N. Donner 7196, 16.viii.1980, 9.67, CHR 411138, AY591853, GU045818. Plagianthus divaricatus J.R.Forst. & G.Forst., New Zealand, Canterbury, Banks Peninsula, Teddington, D. Banks 90/33 and P. Douglass, 16.xi.1990, 9.38, CHR 474070, GU045791, GU045819. Plagianthus regius (Poit.) Hochr., New Zealand, Canterbury, Summit Rd, sign of the Bellbird, R. Elder and J. Thompson, 27.xi.1988, 2.153, CHR 465352, GU045792, GU045820.