A molecular phylogenetic analysis of Diuris (Orchidaceae) based on AFLP and ITS reveals three major clades and a basal species
James O. Indsto A C D , Peter H. Weston A and Mark A. Clements BA National Herbarium of NSW, Mrs Macquaries Road, Sydney, NSW 2000, Australia.
B Centre for Plant Biodiversity Research, National Botanic Gardens, Canberra, ACT 2601, Australia.
C Institute for Conservation Biology, University of Wollongong, NSW 2522, Australia. Present address: Forensic Science Services Branch, Forensic Services Group, NSW Police Force.
D Corresponding author. Email: inds1jam@police.nsw.gov.au
Australian Systematic Botany 22(1) 1-15 https://doi.org/10.1071/SB08029
Submitted: 17 June 2008 Accepted: 24 November 2008 Published: 11 March 2009
Abstract
Diuris is a terrestrial orchid genus of at least 61 and possibly more than 100 species, restricted to Australia except for one species endemic in Timor. Distinctive species groups have respective eastern and western centres of distribution. Although species affinities have been vaguely understood for many years, no formal infrageneric treatment has been undertaken as Diuris possesses few reliable morphological characters for a classification system. We have undertaken cladistic parsimony and Bayesian phylogenetic analyses of Diuris by using the ITS1–5.8S–ITS2 region of nuclear rDNA and morphological characters, with a subset of samples also studied by amplified fragment length polymorphism (AFLP) as an independent test of phylogenetic relationships. Four major clades with strong bootstrap support were resolved and are named here according to a recently published classification; D. sulphurea forms a lineage (subg. Paradiuris) of its own that is well supported as the sister to the rest of Diuris. Two other major eastern clades contained species related to D. maculata (subg. Xanthodiuris) and D. punctata (subg. Diuris), respectively. Although these latter two subgenera are genetically well resolved, there is minimal genetic variation at species level, consistent with recent, rapid speciation. A fourth clade (subg. Hesperodiuris) has a centre of distribution in Western Australia, and has more genetic and morphological variation than the eastern subgenera. Total evidence analysis provides support for the western clade being sister group to the two eastern subgenera Diuris and Xanthodiuris; however, this relationship was not resolved by molecular data. Hybridisation is known to be common among species within subgenera Diuris and Xanthodiuris. Instances of incongruence between different datasets were found suggestive of hybridisation events between species of different sections of Diuris.
Acknowledgements
This work has been undertaken as part completion of a Master of Science degree by JI. Financial support for the project was from a Hermon Slade Orchid Trust grant. The Westmead Millennium Institute (University of Sydney) kindly provided laboratory facilities. John Riley in particular gave much advice on Diuris colony locations and flowering times. Michael Batley provided several Diuris samples used in this study. We also thank Debby McGerty for editing of graphics.
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Appendix 1. Diuris species list compiled by Mark Clements, current as at December 2008 (see http://anbg.gov.au/cpbr/cd-keys/orchidkey/html/currentspecies.html)
Diuris Sm., Trans. Linn. Soc. London 4: 222 (1798). Type species: Diuris aurea Sm.
subgen. Diuris
Sect. Diuris
Synonym: Diuris sect. Flaviflorae G.Don in Loudon’s Hortus Britannicus 368 (1830). Type species: Diuris aurea Sm., fide Jones and Clements (2006).
Diuris aurea Smith, Exotic Bot. 1: 15, t. 9 (1805).
Diuris byronensis D.L.Jones, Orchadian 14(3): 132–133, f. 1, t. (2003).
Diuris chrysantha D.L.Jones et M.A.Clem., Proc. Roy. Soc. Queensland 98: 130–132, f. 7 (1987).
Diuris disposita D.L.Jones, Austral. Orch. Res. 2: 55, f. 69 (1991).
Diuris flavescens D.L.Jones, Austral. Orch. Res. 2: 56, f. 71 (1991).
Diuris luteola D.L.Jones et B.Gray, Austral. Orch. Res. 2: 57–58, f. 73 (1991).
Diuris unica D.L.Jones, Austral. Orch. Res. 5: 82, f. 3.13, t. (21 Dec. 2006).
Diuris secundiflora Fitzg., Austral. Orch. 1(4): [t. 9] (1878).
Diuris tricolor Fitzg., J. Bot. 23: 137 (1885).
sect. Purpureo-albae G.Don in Loudon’s Hortus Britannicus 368 (1830). Type species: Diuris alba R.Br., fide Jones and Clements (2006).
Diuris alba R.Br., Prod. 326 (1810).
Diuris arenaria D.L.Jones, Orchadian 12(12): 567–568, f. 1, t. (1999).
Diuris callitrophila D.L.Jones, Orchadian 14(3): 133–135 (2003).
Diuris curta D.L.Jones, Austral. Orchid. Res. 5: 76–77, f. 3.9 (21 Dec. 2006).
Diuris daltonii (C.Walter) D.L.Jones et M.A.Clem., Orchadian 14(8): Sci. Suppl. xiv (June 2004).
Diuris dendrobioides Fitzg., Austral. Orch. 1(7): [t. 3] (1882).
Diuris fragrantissima D.L.Jones et M.A.Clem., Austral. Orch. Res. 1: 68 (1989).
Diuris minor (Benth.) D.L.Jones et M.A.Clem., Orchadian 14(8): Sci. Suppl. xiv (June 2004).
Diuris oporina D.L.Jones, Austral. Orch. Res. 2: 59–60, f. 77 (1991).
Diuris parvipetala (Dockrill) D.L.Jones et M.A.Clem., Proc. Roy. Soc. Queensland 98: 132 (1987).
Diuris punctata Sm. var. punctata Exotic Bot. 1: 13, t. 8 (1804).
Diuris punctata Sm. var. sulphurea Rupp, Proc. Linn. Soc. New South Wales 69: 73 (1944).
subgen. Xanthodiuris D.L.Jones et M.A.Clem., Orchadian 15(5): 203 (2006). Type species: Diuris maculata Sm.
sect. Xanthodiuris
Diuris aequalis F.Muell. ex Fitzg., Austral. Orch. 1(2): [t. 6] (1876).
Diuris bracteata Fitzg., Austral. Orch. 2(4): [t. 2] (1889).
Diuris cuneilabris Rupp, Proc. Linn. Soc. New South Wales 73: 134, f. 1 (1948).
Diuris goonooensis Rupp, Victorian Naturalist 72: 110 (1955).
Diuris maculata Sm., Exotic Bot. 1: 57, t. 30 (1804–05).
Diuris nigromontana D.L.Jones, Orchadian 15(12): 550–551, t. (June 2008).
Diuris pardina Lindl., Gen. sp. orchid. pl. 507 (1840).
Diuris platichila Fitzg., Austral. Orch. 2(4): [t. 3] (1891).
Diuris semilunulata Messmer in Rupp, Orch. New South Wales 139–140 (1943).
sect. Abbreviatae D.L.Jones et M.A.Clem., Orchadian 15(5): 203–204 (2006). Type species: Diuris abbreviata Benth.
Diuris abbreviata Benth., Fl. Austral. 6: 329 (1873).
Diuris exitela D.L.Jones, Austral. Orch. Res. 2: 55–56, f. 70 (1991).
Diuris praecox D.L.Jones, Austral. Orch. Res. 2: 60, f. 78 (1991).
sect. Pedunculatae D.L.Jones et M.A.Clem., Orchadian 15(5): 204 (2006). Type species: Diuris pedunculata R.Br.
Diuris basaltica D.L.Jones, Austral. Orch. Res. 5: 75–76, f. 3.9 (21 Dec. 2006).
Diuris behrii Schldl., Linnaea 20: 572 (1849).
Diuris chryseopsis D.L.Jones, Austral. Orch. Res. 3: 74–75, f. 4.1 (1998).
Diuris eborensis D.L.Jones, Austral. Orch. Res. 5: 77–78, f. 3.10, t. (21 Dec. 2006).
Diuris fucosa D.L.Jones, Austral. Orch. Res. 5: 78, f. 3.11, t. (21 Dec. 2006).
Diuris gregaria D.L.Jones, Austral. Orch. Res. 5: 79–80, f. 3.12 (21 Dec. 2006).
Diuris lanceolata Lindl., Gen. sp. orchid. pl. 508 (1840).
Diuris monticola D.L.Jones, Austral. Orch. Res. 3: 76–77, f. 4.3 (1998).
Diuris ochroma D.L.Jones, Muelleria 8(2): 182–184, f. 2 d-f (1994).
Diuris pedunculata R.Br., Prod. 316 (1810).
Diuris protena D.L.Jones, Austral. Orch. Res. 5: 81–82 (21 Dec. 2006).
Diuris subalpina D.L.Jones, Orchadian 15(12): 551, t. (June 2008).
Diuris venosa Rupp, Proc. Linn. Soc. New South Wales 51: 313, f. 1–6 (1926).
subgen. Hesperodiuris D.L.Jones et M.A.Clem., Orchadian 15(5): 204 (2006). Type species: Diuris laxiflora Lindl.
sect. Hesperodiuris
Synonym: Diuris subgen. Hesperodiuris sect. Setaceae D.L.Jones et M.A.Clem., Orchadian 15(5): 204 (2006). Type species: Diuris setacea R.Br.
Diuris brevifolia R.S.Rogers, Trans. & Proc. Roy. Soc. South Australia 46: 148 (1922).
Diuris carinata Lindl., Gen. sp. orchid. pl. 510 (Sep. 1840).
Diuris concinna D.L.Jones, Austral. Orch. Res. 2: 53–54, f. 67 (1991).
Diuris drummondii Lindl. in Edwards’s, Bot. Reg. 1–23: Swan Riv. Append. li (1840).
Diuris emarginata R.Br., Prod. 316 (1810).
Diuris filifolia Lindl. in Edwards’s, Bot. Reg. 1–23: Swan Riv. Append. li (1840).
Diuris heberlei D.L.Jones, Austral. Orch. Res. 2: 56–576, f. 72 (1991).
Diuris immaculata D.L.Jones, Austral. Orch. Res. 5: 80–81 (21 Dec. 2006).
Diuris laxiflora Lindl. in Edwards’s, Bot. Reg. 1–23: Swan Riv. Append li (1840).D.
Diuris micrantha D.L.Jones, Austral. Orch. Res. 2: 58–59, f. 75 (1991).
Diuris pauciflora R.Br., Prod. 316 (1810).
Diuris picta J.Drummond in Hooker’s, J. Bot. 5: 347 (1853).
Diuris setacea R.Br., Prod. 316 (1810).
sect. Pyrophilae D.L.Jones et M.A.Clem., Orchadian 15(5): 204 (2006). Type species: Diuris laevis Fitzg.
Diuris laevis Fitzg., Gard. Chron. (new ser.), 17: 495 (1882).
Diuris purdiei Diels, J. Proc. Mueller Bot. Soc. Western Australia_1(11): 79 (1903).
sect. Palustres D.L.Jones et M.A.Clem., Orchadian 15(5): 204–205 (2006). Type species: Diuris palustris Lindl.
Diuris palustris Lindl., Gen. sp. orchid. pl. 507 (1840).
sect. Suffusae D.L.Jones et M.A.Clem., Orchadian 15(5): 205 (2006). Type species: Diuris longifolia R.Br.
Diuris amplissima D.L.Jones, Austral. Orch. Res. 2: 53–54, f. 65 (1991).
Diuris brumalis D.L.Jones, Austral. Orch. Res. 2: 53, f. 66 (1991).
Diuris conspicillata D.L.Jones, Austral. Orch. Res. 2: 54–55, f. 68 (1991).
Diuris corymbosa Lindl. in Edwards’s, Bot. Reg. 1–23: Swan Riv. Append. li (1840).
Diuris longifolia R.Br., Prod. 316 (1810).
Diuris magnifica D.L.Jones, Austral. Orch. Res. 2: 58, f. 74 (1991).
Diuris orientis D.L.Jones, Austral. Orch. Res. 3: 77–78, f. 4.4 (1998).
Diuris porrifolia Lindl. in Edwards’s, Bot. Reg. 1–23: Swan. Riv. Append. li (1840).
Diuris pulchella D.L.Jones, Austral. Orch. Res. 2: 61, f. 79 (1991).
Diuris recurva D.L.Jones, Austral. Orch. Res. 2: 61–62, f. 80 (1991).
subgen. Paradiuris D.L.Jones et M.A.Clem., Orchadian 15(5): 205 (2006). Type species: Diuris sulphurea R.Br.
Diuris sulphurea R.Br., Prod. 316 (1810).
subgen. Timordiuris D.L.Jones et M.A.Clem., Orchadian 15(5): 205 (2006). Type species: Diuris fryana Ridl.
Diuris fryana Ridl. in H.O.Forbes, Nat. Wand. East. Archip. 519 (1885).
Natural hybrids
Diuris × fastidiosa R.S.Rogers, Trans. & Proc. Roy. Soc. South Australia 51: 6–7 (1927).
Diuris × nebulosa D.L.Jones, Austral. Orch. Res. 2: 59, f. 76 (1991).
Diuris × palachila R.S. Rogers, Trans. & Proc. Roy. Soc. South Australia 31: 209–210 (1907).
Diuris × polymorpha Messmer in Rupp, Orchids New South Wales, Suppl. 142 (1943).