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Morphological and Molecular Data Reveal Three Rather Than'one Species

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Morphological and Molecular Data Reveal Three Rather Than'one Species CSIRO PUBLISHING Australian Systematic Botany, 2012, 25, 188–201 http://dx.doi.org/10.1071/SB11032 Morphological and molecular data reveal three rather than one species of Sicyos (Cucurbitaceae) in Australia, New Zealand and Islands of the South West Pacific Ian R. H. Telford A,D, Patrizia Sebastian B, Peter J. de Lange C, Jeremy J. Bruhl A and Susanne S. Renner B AN.C.W. Beadle Herbarium, University of New England, Armidale, NSW 2351, Australia. BSystematic Botany and Mycology, University of Munich, Menzinger Strasse 67, 80638, Munich, Germany. CEcosystems and Species Unit, Department of Conservation, Private Bag 68908, Newton, Auckland 1145, New Zealand. DCorresponding author. Email: [email protected] Abstract. Morphometric data have long suggested the existence of three species of Sicyos in Australia and New Zealand. Molecular data now corroborate this and place the three species in the context of the remaining 72 species of Sicyos, most of them in the New World. We here describe and illustrate Sicyos undara I.Telford & P.Sebastian from northern Queensland and S. mawhai I.Telford & P.Sebastian from New Zealand, and emend the circumscription of S. australis to account for the removal of these two extraneous elements. We also provide a key to the three species, map their ranges, and note their habitat and conservation status. Received 26 August 2011, accepted 7 December 2011, published online 6 June 2012 Introduction specimens from Gippsland, Victoria, and held in MEL as Sicyos L. is a genus of 75 species, most of them occurring in S. rivularis MS, a manuscript name never validated. Central and South America, although the genus also underwent Doubts about Hooker’s broad species concept were first radiation on Hawaii and has two endemic species on the expressed by Allan (1961) in the ‘Flora of New Zealand’ who Galapagos Islands (Sebastian et al. 2012). In the South West noted that the South West Pacific collections needed critical Pacific region, Sicyos has been recorded from eastern Australia, comparison with American material; still, he decided to Tasmania, New Zealand and smaller islands associated to these maintain the name S. angulatus for New Zealand indigenous major landmasses, as well as Lord Howe Island, Norfolk Island plants. Telford (1982), in his treatment of the Cucurbitaceae for and the Kermadec Islands. Molecular phylogenies for the the ‘Flora of Australia,’ took up Endlicher’s name S. australis Cucurbitaceae show that Sicyos and 11 other genera (with for Australian specimens and also stressed the heterogeneity of some 265 species altogether) form a clade in which the the South West Pacific material and the need for a revision. monotypic Queensland genus Nothoalsomitra I.Telford is Comparing S. angulatus material from New Zealand and sister to the remaining genera. This newly circumscribed tribe, America, Sykes (in Connor and Edgar 1987) found clear named Sicyoeae, includes several New World genera, and also differences, but still referred the New Zealand material to the large Old World genus Trichosanthes L. and the pantropical S. australis, a decision followed by other New Zealand authors genus Luffa Mill. (Schaefer and Renner 2011). (Webb et al. 1988; de Lange et al. 2004, 2006, 2009b). For the Australian material of Sicyos, two names are available, Chromosome numbers (de Lange and Murray 2002) and namely S. australis Endl. (Endlicher 1833) and S. fretensis Hook. observations of morphological differences (Brandon et al. f. (Hooker 1847). Both appear on specimens housed at K and BM, 2004; Forester and Townsend 2004) in the meantime have but have rarely been used in Australian herbaria. Hooker (1860) made it clear that more than one species of Sicyos occurs in later subsumed his Australian species, together with New Zealand New Zealand. Post-1982 collections from Undara Lava Tubes material, into a broadly defined S. angulatus L., a species based on in northern Queensland also suggested the existence of a new material from the eastern United States. Probably because of species in Australia, prompting a phenetic analysis of 34 Hooker’s eminent status and the scarcity of material in European herbarium sheets, covering the morphological and geographic herbaria, this broad concept became widely accepted (Bentham range of material available by 1997 (I. R. H. Telford, unpubl. 1866; Bailey 1900; Allan 1961; Beadle et al. 1962; Curtis 1963; data). The results strongly supported the existence of three Willis 1972; Beadle 1976). Ferdinand von Mueller had annotated species in the South West Pacific region, a finding supported Journal compilation Ó CSIRO 2012 www.publish.csiro.au/journals/asb New species of Sicyos in the South West Pacific Australian Systematic Botany 189 by a genus-wide phylogeny (Sebastian et al. 2012). Here, we characters, and rehydrated material for floral characters describe, map and illustrate the two new species, S. undara and (Appendix 1); vegetative characters, such as leaf shape and S. mawhai (names used hereafter in the present paper), pubescence, were avoided because of their well known and provide an emended circumscription of S. australis sensu plasticity in the family. stricto. We also provide a key to all three species. The data matrix was analysed using the pattern analysis software package PATN version 3.03 (Belbin and Collins Materials and methods 2006), with all characters given equal weight and range- Phenetic analysis standardised by using the Gower metric association measure. Two options of PATN (Belbin 1990a, 1990b) were used to define We here use a morphological species concept based on suites population groupings, namely, cluster analysis, employing the of covarying attributes (Stuessy 1990), and we tested the flexible unweighted pair group arithmetic averaging (UPGMA), delimitation of our new species using graphical and statistical presented graphically as a phenogram, and ordination analysis, analyses of morphological datasets (Sneath and Sokal 1973). The using semi-strong hybrid multidimensional scaling (SSH MDS) morphological dataset was derived from herbarium specimens in a three-dimensional scatter plot. held at AK, BM, BRI, CANB, CHR, HO, JCU, K, MEL, NE, NSW and W. Each operational taxonomic unit (OTU) consisted of a single gathering, whether mounted on a single sheet or Molecular analysis multiple sheets. Thirty-four OTUs were selected to cover the For the molecular phylogeny shown here, we used a modified range of the genus in Australia and New Zealand (Table 1), and version of the matrix used in Sebastian et al.(2012). That study the characters used are listed in Table 2. States were assigned had 112 accessions, representing 87 species of Sicyoeae, using dried herbarium specimens for inflorescence, fruit and seed including the type species of all relevant generic names. Here, Table 1. Operational taxonomic unit (OTU) numbers, codes used in analyses and vouchers for phenetic analysis Voucher data present collector’s name, number and herbarium where housed. Herbarium accession number is used when collector’s number is absent. Where more than one collector involved, only the first collector is given. Herbarium codes follow Index Herbariorum, available at http://sweetgum.nybg.org/ih/ (accessed 12 October 2011) OTU Code Locality Collector 1 NZ1 Cuvier Island, New Zealand R. Moynihan AK119648 (AK) 2 NZ2 Hen Island, New Zealand D. Merton CHR183501(CHR) 3 NZ3 Little Barrier Island, New Zealand W.R. Sykes 147/85 (CHR) 4 NZ4 Auckland, New Zealand J.J. Bruhl 2915 (NE) 5 NZ5 Coromandel Island, New Zealand A.E. Esler 3584 (CHR) 6 NZ6 Tahuna Road, Te Teko, New Zealand W. Stahel CHR566019 (CHR) 7 NZ7 White Pine Bush, New Zealand M. Steverinson CHR551181(CHR) 8 NZ8 Poor Knights Island, New Zealand G.N. Park CHR276071 (CHR) 9 NZ9 Pakaraka, New Zealand K. Riddell AK238904 (AK) 10 NZ10 Whenuakite, New Zealand C. Wallace AK199785 (AK) 11 NZ11 Three Kings Islands, New Zealand A.E. Wright 5266 (AK) 12 NZ12 Three Kings Islands, New Zealand A.E. Wright 6061 (AK) 13 KI1 Kermadec Islands T.F. Cheeseman MEL591390 (MEL) 14 KI2 Kermadec Islands C.J. West CHR518213 (CHR) 15 KI3 Kermadec Islands W. Sykes 1523/K (CHR) 16 NI1 Norfolk Island W. Robinson (K) 17 NI2 Norfolk Island F. Bauer 110 (W) 18 LH1 Lord Howe Island leg. ign. MEL593267–8 (MEL) 19 LH2 Lord Howe Island J. Fullagar 118 (MEL) 20 LH3 Lord Howe Island leg. ign. (K) 21 TS1 Sisters Island, Tasmania J. Whinray 57 (CANB) 22 TS2 Sisters Island, Tasmania J. Whinray 492 (CANB) 23 VG1 Buchan River, Victoria F. Mueller MEL593278(MEL) 24 VG2 Tambo River, Victoria F. Mueller MEL593283 (MEL) 25 NS1 Tuross River, New South Wales J. Whaite 478 (NE) 26 NS2 Cambewarra Mountain, New South Wales I.R. Telford 11793 (CANB) 27 NS3 Nymboida River, New South Wales J.J. Bruhl 2102 (NE) 28 QL1 Killarney, Queensland I.R. Telford 12909 (NE) 29 QL2 Isla Gorge, Queensland P. Sharpe 660 (BRI) 30 QL3 Eungella, Queensland N. Byrnes 3699 (BRI) 31 QL4 Fanning River Caves, Queensland B. Jackes S8643 (JCU) 32 QL5 Wind Tunnel, Undara, Queensland I.R. Telford 13319(NE) 33 QL6 Pinwill Cave, Undara, Queensland V.J. Neldner 2780 (BRI) 34 QL7 Pinwill Cave, Undara, Queensland I.R. Telford 13317 (NE) 190 Australian Systematic Botany I. R. H. Telford et al. Table 2. Morphological characters used for ordination and cluster The matrix comprised six chloroplast regions, the trnL intron, analysis of Sicyos in the South West Pacific the trnL–F, rpl20–rps12, trnS–G and psbA–trnH intergenic Number Character spacers, and the rbcL gene, and the nuclear rDNA internal transcribed spacers ITS1 and ITS2, with the intervening 5.8 S Female inflorescence gene, for a length of 5399 aligned positions (4527 chloroplast 1 Peduncle length (mm) and 872 nuclear). GenBank accession numbers and voucher fl 2 Flowers per in orescence information are listed in table S1 of Sebastian et al.(2012).
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