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Home , Acacia harpophylla, Entelea, Lachnagrostis, Myoporum laetum, Sicyos

CSIRO PUBLISHING Australian Systematic , 2012, 25, 188–201 http://dx.doi.org/10.1071/SB11032

Morphological and molecular data reveal three rather than one species of Sicyos () in , 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 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, , and held in MEL as Sicyos L. is a of 75 species, most of them occurring in S. rivularis MS, a manuscript name never validated. Central and , 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 ‘’ 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 , New Zealand and smaller islands associated to these maintain the name S. angulatus for New Zealand indigenous major landmasses, as well as , . Telford (1982), in his treatment of the Cucurbitaceae for and the . Molecular phylogenies for the the ‘,’ 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). had annotated species in the South West Pacific region, a finding supported

Journal compilation 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, 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). 3 Corolla diameter (mm) DNA extraction, polymerase chain reactions, sequencing and Fruit 4 Fruit length (mm) data preparation were carried out using standard approaches 5 Fruit diameter (mm) (Sebastian et al. 2010a, 2012). Maximum likelihood (ML) 6 Aculei length (mm) analyses and bootstrap searches using 500 replicates were 7 Aculei per fruit carried out using RAxML version 7.2.8 (Stamatakis 2006), 8 Seed length (mm) relying on the GTR + G model, with model parameters Male inflorescence estimated over the duration of specified runs. 9Inflorescence length (mm) 10 Peduncle length (mm) 11 Flower number per inflorescence Results 12 Pedicel length (mm) Phenetic analysis 13 Corolla diameter (mm) Staminal column length (mm) The 34 specimens from New Zealand, Tasmania, Lord Howe 15 Staminal head diameter (mm) Island, Kermadec Islands, Norfolk Island, and the Australian states Victoria, New South Wales and Queensland used in the phenetic analyses clearly fall into three groups on the basis of we focus the sample to 34 of these to provide two accessions for UPGMA clustering (Fig. 1) and SSH MDS ordination (Fig. 2). each Australian and New Zealand species of Sicyos, and sound Table 3 shows the correlation of characters with ordination sampling across other relevant species. patterns: Fruit and seed attributes contributed most to phenetic 0.066 5 0.232 3 0.398 2 0.564 0 0.729 9

NZ1 NZ2 KI2 NZ3 NZ5 KI1 Sicyos mawhai NZ4 NZ8 KI3 NZ11 NZ12 NZ6 NZ9 NZ7 NS1 NI2 VG2 LH3 LH2 VG1 Sicyos australis QL2 LH1 TS2 TS1 NZ10 NS3 QL1 NI1 NS2 QL3 QL4 Sicyos undara QL5 QL6 QL7

Fig. 1. Phenogram based on Gower association and flexible unweighted pair group arithmetic averaging (UPGMA) of the operational taxonomic units (OTUs) of Sicyos in the South West Pacific. See Table 1 for OTU coding and vouchers. New species of Sicyos in the South West Pacific Australian Systematic Botany 191

(judging from branch lengths of other species-level groups in Sicyos mawhai Sicyos) and, moreover, shows Australian S. undara as sister to S. australis and the New Zealand endemic S. mawhai. These three species stem from a common ancestor (Fig. 3) and are sister to a Sicyos acstralis clade from the South Western United States, Mexico and Bolivia. The American species S. angulatus is placed distantly from 1.543 0.875 the Australian–New Zealand clade (Fig. 3), matching the morphological differences observed by Sykes in Connor and Edgar (1987; see also Webb et al. 1988). Sicyos undara Discussion Many Cucurbitaceae are successful transoceanic dispersers fi 0.781 (Schaefer et al. 2009), and Sicyos, with four trans-Paci c –0.925 range disjunctions (Sebastian et al. 2012), constitutes a prime example of this. The latter study dated the trans-Pacific dispersal of the ancestor of the Australian–New Zealand clade of Sicyos to – –1.194 1.416 5.6 1.7 million years (Ma) ago, the divergence of the Queensland species S. undara to 3.6–0.5 Ma, and the arrival of the ancestor of Fig. 2. Three-dimensional ordination of semi-strong hybrid S. mawhai in New Zealand to 1.6–0.1 Ma. S. australis arrived in multidimensional scaling (SSH MDS) for organisational taxonomic units New Zealand more recently. (OTUs) of Sicyos in the South West Pacific. Stress = 0.0847. Dispersal across the ~2000 km of ocean between Australia and New Zealand has been inferred for many other groups (Perrie et al. 2003; Chandler et al. 2007; de Lange et al. 2010; Tay Table 3. Correlation of character values with correlation of attributes for semi-strong hybrid multidimensional scaling (SSH MDS) ordination et al. 2010; Prebble et al. 2011) and numerous species, including of Sicyos in the South West Pacific apparently recent arrivals (Sykes and de Lange 1993; de Lange et al. 2009a), show trans-Tasman disjunctions. Character XYZR2 As is true of their closest New World relatives, the small, fi 5 0.798 0.355 –0.488 0.923 one-seeded fruits of the three South West Paci c species bear 8 0.298 0.666 –0.684 0.918 retrorsely barbed spines, which may facilitate transport in birds’ 7 0.255 0.377 –0.890 0.821 plumage, and both S. australis and S. mawhai sometimes occur in 15 –0.176 0.310 –0.934 0.810 coastal vegetation where they may come in contact with petrels or 13 0.662 0.616 –0.428 0.796 shearwaters, which regularly travel between the southern Pacific 4 0.467 0.623 –0.627 0.782 landmasses. 12 0.788 0.616 –0.019 0.734 Sicyos mawhai is the first endemic New Zealand species of – – 9 0.557 0.171 0.812 0.710 the Cucurbitaceae family known. Both Sicyos species now 3 0.800 0.009 –0.601 0.707 known from New Zealand are threatened by cucurbit diseases, 14 –0.628 0.123 –0.769 0.625 6 –0.649 –0.186 –0.738 0.620 such as cucumber mosaic virus (CMV) and zucchini yellow 10 0.292 –0.457 –0.840 0.534 mosaic virus (ZYMV) (Delmiglio and Pearson 2006), which 11 0.966 –0.023 –0.257 0.508 may also be responsible for the decline of the two Sicyos 1 0.778 0.187 0.600 0.446 species on the Galápagos archipelago (Sebastian et al. 2010b). 2 0.463 0.797 –0.388 0.265 Australian populations, by contrast, appear healthy, with no obvious decline through disease (I. R. H. Telford, pers. obs.). differences. The results indicate that two species should be recognised in Australia, the widespread S. australis (OTUs 16–31) and an entity from Undara (OTUs 32–34), which we Sicyos australis Endl., Prodr. Fl. Norfolk. 67 (1833) emend. describe below. S. fretensis (OTUs 21, 22) and Mueller’s I.Telford unpublished S. rivularis (OTUs 23, 24) are grouped within Type: NORFOLK ISLAND, western mountain, F. Bauer 110; S. australis. New Zealand material groups into two species, holo: W!. the putative new species (OTUs 1–5, 8, 9, 11–16), named ’ S. mawhai below, and those (OTUs 6, 7, 10) grouping with S. fretensis Hook.f., Hooker s London J. Bot. 6: 473 bis (1847). S. australis, confirming their placement by Brandon et al. Type: Sisters’ Island, E coast of Flinders Island, R. Gunn [Oct., (2004). 1844, J. Milligan 563]; holo: K!; iso: BM, HO, K!, MEL!. S. angulatus auct. non L.: Hooker, 143 (1856); Bentham, 3: 322 Molecular analysis (1866); Bailey, 701 (1900); Beadle et al., 199 (1963); Curtis, 237 The interpretation of the phenetic analyses, as indicating the (1963); Willis, 2: 264 (1972); Beadle, 271 (1976). existence of three biological species, matches the molecular S. aff. australis (b) (AK 289786; Mangere Stonefields) de Lange et al. phylogeny (Fig. 3), which recovers three species-level groups (2009b). 192 Australian Systematic Botany I. R. H. Telford et al.

Fig. 3. Maximum likelihood phylogram for 33 taxa of Sicyoeae based on combined sequences from chloroplast and nuclear data (5399 aligned nucleotides), analysed under the GTR + G substitution model. The tree is rooted on Linnaeosicyos. The Australasian clade is highlighted. Likelihood bootstrap values 75% are given at the nodes; geographic origin of the sequenced specimens follows the species name and asterisks indicate generic type species.

Monoecious herbaceous climber; stems annual, up to 10 m densely echinulate with barbed aculei; hypanthium above the long, 5-mm diam., sparsely hirsute with simple multicellular constriction broadly campanulate, ~1.5 mm in diam., minutely hairs and glandular hairs, glabrescent. Tendrils 3–5-branched. glandular hairy surrounding the disc; calyx lobes linear, Leaves: petiole 10–65 mm long, scaberulous; lamina ovate or 0.4–0.7 mm long; corolla 2.5–3.5-mm diam., mostly glabrous broadly ovate in outline, 35–195 mm long, 45–200 mm wide, abaxially, the lobe apices puberulous, glabrous adaxially, white, cordate with the basal sinus broad, the lobes not overlapping, 5-lobed, the lobes triangular–ovate, obtuse, ~1.6 mm long; disc acuminate, shallowly to deeply palmately 5- or 7-lobed, the ~1 mm in diam.; style ~1.4 mm long; stigma 2-lobed, the lobes broadly triangular, acute or acuminate, margins dentate lobes ~0.3 mm long, recurved. Fruit ovate, rarely fusiform, with apiculate teeth, sparsely scaberulous adaxially and 6.4–9.5 mm long, 2.8–4 mm in diam., apically attenuate, the abaxially with simple hairs, more densely and coarsely surface Æglabrous or scabridulous with short hairs, echinate; along veins. Male inflorescence an 8–19-flowered raceme aculei dense, 1.6–3.6 mm long, retrorsely barbed. Seeds 25–110(–155) mm long; peduncle 15(–125) mm long, sparsely ellipsoidal, 4–5.5 mm long, 2.8–3.4 mm wide, brown. scaberulous; rhachis glandular hairy. Male flowers: pedicels 3–11 mm long; hypanthium broadly campanulate, 2.4–2.7-mm diam.; calyx lobes linear, 0.4–0.7 mm long; corolla rotate, Distribution 4.5–7.8 mm in diam., mostly glabrous abaxially, the lobe In eastern Australia, S. australis occurs from Fanning River, apices puberulous, glabrous adaxially, white, 5-lobed, the ~60 km south-west of Townsville, Queensland, south to Orbost, lobes broadly triangular–ovate, obtuse; 2–2.6 mm long; disc Victoria, inland to the Great Dividing Range, and on Sisters’ ~1.2 mm in diam.; staminal column 1.5–1.8 mm long; staminal Island, off Flinders Island, Tasmania (Fig. 4). head 1.7–2.2-mm diam. Female inflorescence an 8–11-flowered In New Zealand, the first collection of which we are aware head; peduncle 8–23(–33) mm long. Female flowers: subsessile; was from Thames, North Island, by T. F. Cheeseman of the ovate, attenuate, ~3 mm long, 1.4-mm diam., minutely and Auckland Museum. The collection date was not recorded; New species of Sicyos in the South West Pacific Australian Systematic Botany 193

150°0′0′′E 160°0′0′′E 170°0′0′′E 10°0′0′′S

20°0′0′′S 20°0′0′′S

30°0′0′′S 30°0′0′′S

40°0′0′′S 40°0′0′′S

km

150°0′0′′E 160°0′0′′E 170°0′0′′E

Fig. 4. Distribution of Sicyos in South-West Pacific. * S. australis; $ S. mawhai; ~ S. undara. Note that this represents the historical range of species; localities where S. australis and S. mawhai are known to be extinct are shown.

however, consultation of the Cheeseman notebooks suggests In New Zealand, S. australis is a species of disturbed some time in the 1870s and 1880s (E. K. Cameron, pers. forest, forest margins, agricultural land, associated wasteland comm.). Recent collections (1988+) on the North Island have and roadsides. In forested habitats, it often associates with been made from Northland, Auckland, Waikato and the Bay of members of the Kunzea ericoides agg., Melicytus ramiflorus, Plenty. Pseudopanax crassifolius, Vitex lucens, Corynocarpus Previously recorded from the Lord Howe and Norfolk Island laevigatus, Dysoxylum spectabile and Metrosideros excelsa. groups, the latter being the type locality of the species, S. australis Otherwise, it has been collected threaded through the exotic has not been collected from either of these island groups for more grass Cortaderia selloana, or climbing over such naturalised than a century and is presumed extinct in these archipelagos shrubs and trees as Ligustrum sinense, L. vulgare, Robinia (Green 1994). pseudacacia, Populus nigra and Solanum mauritianum in wasteland, on occasion as a weed in maize (Zea mays) crops, or growing on ancient basalt stonewalls within the site of a Habitat and ecology former, major pre-European settlement Maori agricultural site. In mainland Australia, S. australis grows in dry rainforest At many of these habitats, S. australis exhibits a ‘weedy’ and (deciduous vine thicket) with Brachychiton australis on basalt invasive nature, which has lead people to regard it as a recent or limestone in northern Queensland, and with B. rupestris naturalisation rather than an indigenous species. and Acacia harpophylla on cracking clay derived from basalt Although the species was first collected from New Zealand in central Queensland. In coastal and near-coastal montane by Cheeseman in the late 19th century, it was not then recorded habitats from Eungella, Queensland, south to Gippsland, again from that country until 1988 (when it was gathered Victoria, S. australis grows in wetter eucalypt forest or on from a remote area of coastal forest on the eastern side of the rainforest margins, at disturbed sites in closed forest, often at Coromandel Peninsula). On this evidence, we think that sheltered sites in gorges and on a wide variety of substrates. S. australis has naturally colonised New Zealand, probably via 194 Australian Systematic Botany I. R. H. Telford et al. trans-Tasman avian dispersal, possibly on several occasions, or Sicyos australis sensu stricto in de Lange et al.(2009b) refers it has persisted undetected in the less well explored parts of the to plants on the Kermadec Islands, then thought to be conspecific northern North Island of New Zealand until recently. Some with that species, but herein shown to be S. mawhai, and the threat evidence suggests that S. australis is much more likely to have assessment therein must lapse. recently colonised New Zealand rather than recently spread from long-established indigenous populations. Notably, there Notes fi ‘ ’ fi is its speci c association at two sites, Jacks Bush ( rst The protologue of S. fretensis attributes the type collection to collected 1993) and Whenuakite (first collected 1988), with Ronald Gunn. Gunn’s private herbarium included specimens another naturally recently established trans-Tasman species, collected by others, sometimes with label data transcribed fl fi Plectranthus parvi orus, also rst recognised from New without collector, and these were often assumed to have been Zealand in 1988 (Sykes and de Lange 1993), whereas several collected by Gunn himself (Buchanan 1988). other Northland occurrences of Sicyos are in the general vicinity of another trans-Tasman colonist recently accepted Illustrations as indigenous to New Zealand, namely, Rorippa laciniata D. L. Jones & B. Gray, Australian Climbing Plants, fig. 227 (de Lange et al. 2009a). Further, on the basis of herbarium (1977); N. G. Walsh & T. J. Entwisle (Eds), Fl. Victoria 3: 384, evidence, it is significant that the first gatherings of fig. 79h (1996); L. Forester & A. Townsend, Threatened Plants of S. australis were all made from remote indigenous forest Northland Conservancy 61 (2004); upper image. remnants in Northland and Coromandel at sites located well away from the main ports of entry to New Zealand, but Additional specimens examined (selection) corresponding to well known sites of trans-Tasman dispersal by vagrants and at times subsequent colonisation and spread NORFOLK ISLAND: 1898, W. Robinson s.n. (K, NSW). NEW (de Lange 1997; Heenan and Forester 1997; de Lange and ZEALAND: North Island: Pakaraka, end of Ludbrooke Road, Pouerua Bush, 28 Apr. 1999, K. Riddell (AK); SW of Pakaraka, edge Norton 1998; de Lange et al. 2011). Once established at these of ‘Jack’s Bush’, S slopes of Pouerua, P.J. Bellingham 591 (AK); sites, it is only to be expected that Sicyos, as an undeniably Mangere, Ihumatao, Otuataua Stonefield, E.K. Cameron 11066 (AK); ‘weedy’ species with fruits well suited to avian and mammalian Coromandel, Whenuakite, off Boat Harbour Road, 10 June 2002, dispersal, would be rapidly spread by a host of indigenous C. Delmiglio (AK); Te Teko, Tahuna Road, Urupa, P.J. de Lange and exotic birds, and widespread mammals such as opossum 6441 & P.B. Cashmore (AK, NSW); White Pine Bush near Awakeri, (Trichosurus vulpecula), let alone by humans. This would maize paddock at end of Ernest Road, 26 May 2006, M. Steverinson & explain its more recent spread throughout wasteland and P.B. Cashmore (AK, CHR, NZFRI). AUSTRALIA: Queensland: North roadsides as well as its ‘spot’ occurrences as a volunteer in Kennedy District: Fanning River, N of Mingela, B.R. Jackes S8643 (JCU, agricultural crops. In this regard, S. australis is following a NSW); South Kennedy District: Eungella, N. Byrnes 3699 & J. Clarkson pattern already typical of many other undeniably long- (BRI); Leichhardt District: Ka Ka Mundi section, Carnarvon National Park, on fire trail from Salvator Rosa, M.B.Thomas 3781 & C.Elmes established indigenous New Zealand plants with the same (BRI); Darling Downs District: Great Dividing Range, ~15 km ENE of dispersal tendencies and which are now common in such Killarney, I.R. Telford 12909, J.J. Bruhl & L.M. Copeland (CANB, M, habitats, e.g. Alternanthera nahui, Epilobium cinereum, NE). New South Wales: Lord Howe Island: no precise loc., J. Fullagar Lachnagrostis filiformis (Heenan et al. 2009). 118 (MEL); North Coast: Nymboi–Binderay National Park, Platypus Flat, J.J. Bruhl 2102 & F.C. Quinn (BRI, CANB, L, MO, NE); Central Phenology Coast: Cambewarra Mountain, Beaumont, I.R. Telford 11793 (AK, CANB, NSW); South Coast: Morton National Park, Sentrybox Flowers and fruits mainly November–July. Canyon, I.R. Telford 10805 (CANB, K, NSW); Tuross River, J. Whaite 478 (NE). Victoria: Buchan River, F. Mueller s.n. MEL Chromosome number 593278 (MEL). Tasmania: Sisters’ Island, J. Whinray 500 (CANB, 2n = ~24, from New Zealand material (de Lange and Murray MEL). CULTIVATED: ex New Zealand, Northland, Waiomio 2002). Stream, Waiomio Caves, Kawiti property, P.J. de Lange 5262 (AK).

Conservation status Sicyos undara I.Telford & P.Sebastian, sp. nov. In mainland Australia, S. australis is widespread and common Ab affini Sicyo australi inflorescentiis foemineis paucifloris throughout its range and is not considered at risk. It is also longius et tenuius pedunculatis nec non fructibus minoribus conserved in many national parks, including Eungella, Isla parcius et minutius aculeatis differt. Gorge, Main Range, Lamington and Springbrook in Type: AUSTRALIA: Queensland: Cook District: Undara Queensland, Border Ranges, Namoi-Binderay, Dorrigo, Oxley Volcanic National Park, 50 m W of Pinwill Cave entrance, 27 Wild Rivers and Morton in New South Wales. March 2009, I.R. Telford 13317 & P. Sebastian; holo: BRI; iso: In New Zealand, S. australis is given a threat status (under the BISH, CANB, CHR, CNS, K, M, MO, NE, US (Fig. 5: isotype informal tag or phrase name S. aff. australis (b) (AK 289786; NE). Mangere Stonefields) of ‘Non-resident native/coloniser’ Monoecious herbaceous climber; stems annual, up to 5 m (Townsend et al. 2008) in de Lange et al.(2009b). Although long, up to 2.5-mm diam., sparsely hirsute with simple this category predates recognition of the collection by Cheeseman multicellular hairs and glandular hairs, densely so at nodes, cited above, it probably remains valid because extinction and glabrescent. Tendrils 3- or 4-branched. Leaves: petiole recolonisation appear more likely than a long-lived seed bank. 10–35 mm long, sparsely pilose with multicellular and glandular New species of Sicyos in the South West Pacific Australian Systematic Botany 195

Fig. 5. Isotype of Sicyos undara I.Telford & P.Sebastian. hairs; lamina subtriangular or broadly ovate in outline, shallowly 3- or 5-lobed, the lobes triangular, acuminate, 20–117 mm long, 20–94 mm wide, cordate with the basal margins dentate with apiculate teeth, hispid on both surfaces, sinus broad, the lobes not overlapping, acuminate, palmately more coarsely so on veins abaxially. Male inflorescence a 196 Australian Systematic Botany I. R. H. Telford et al.

7–25-flowered raceme, 35–155(–180) mm long; peduncle Tunnel, 23 March 1994, D. Hansman s.n. (CANB); Undara Volcanic 24–108 mm long, glabrous; rhachis glandular hairy. Male National Park, Wind Tunnel complex, I.R. Telford 13319 & P. Sebastian flowers: pedicel 5–21 mm long; hypanthium broadly (BRI, CANB, CNS, M, NE). campanulate, ~1.8 mm in diam.; calyx lobes linear, ~0.7 mm long; corolla rotate, 3–5.7 mm in diam., abaxially and Sicyos mawhai I.Telford & P.Sebastian, sp. nov. adaxially glabrous, the margins glandular hairy, greenish Ab affini Sicyo australi inflorescentiis floribusque masculis white, 5-lobed, the lobes triangular; 1–1.8 mm long; disc majoribus, floribus foemineis majoribus in quoque capitulo ~1 mm in diam.; staminal column 0.9–1.1 mm long; staminal numerosioribus nec non aculeis fructuum longioribus discrepat. head 0.7–0.8 mm in diam. Female inflorescence a 4–8-flowered Type: NEW ZEALAND: North Island: Cuvier Island, large head; peduncle (20–)32–74 mm long. Female flowers: subsessile; patch scrambling over bracken on ridgetop, West Point, 25 May ovary ovate, ~1.2 mm long, ~0.7 mm in diam., echinulate with 1980, A.E. Wright 3583; holo:. AK!; iso: A, AD!, AKU, BISH!, barbed aculei; hypanthium above the constriction broadly CM, LTU, NA, WELT. campanulate, ~1.3 mm in diam.; calyx lobes linear, ~0.7 mm long; corolla 2.6–3 mm in diam., glabrous abaxially and S. angulatus auct. non L.: Hooker, 72 (1852); Laing, 38 (1915); Allan, adaxially, the margins minutely glandular papillose, greenish- 319 (1961). white, 5-lobed, lobes triangular, ~1 mm long; disc ~0.5 mm in S. australis auct. non Endl.: Sykes in Webb et al.(1988); de Lange et al. diam; style ~1.5 mm long; stigma 2-branched, the branches (2009b). recurved. Fruit ovate or subglobose, 4–5.8 mm long, S. aff. australis (a) (AK 252822): de Lange et al.(2009b). 2.2–2.6 mm in diam., apically rounded or obtuse, the surface glabrous, echinate; aculei scattered, 1.4–2.8 mm long, retrorsely Monoecious herbaceous climber; stems annual, up to 10 m barbed. Seeds Æovate, ~3.5 mm long, 2 mm wide, cream (Fig. 5). long, up to 2.5 mm in diam., sparsely hirsute with simple multicellular celled hairs and glandular hairs, glabrescent. – – – Distribution Tendrils 3 5-branched. Leaves: petiole 25 75( 135) mm long, minutely glandular hairy; lamina broadly ovate in outline, Sicyos undara is known only from the Undara Lava Tubes east of 45–135 mm long, 60–165 mm wide, cordate with the basal Mount Surprise, Queensland (Fig. 4). sinus Æclosed by overlapping lobes, acuminate, shallowly palmately 5- or 7-lobed, the lobes rounded–triangular, margins Habitat and ecology coarsely dentate with apiculate teeth. Male inflorescence a Sicyos undara grows in skeletal clay loam in boulder gullies 14–35-flowered raceme (45–)70–85(–205) mm long; peduncle formed by collapsed lava tubes. The vegetation is deciduous (35–)55–90(–125) mm long. Male flowers: pedicels vine thicket (dry rainforest) dominated by Brachychiton (3.5–)6–12(–27) mm long; hypanthium broadly campanulate, australis, Ficus virens and Melia azederach, with Pittosporum 3.8–4.2-mm diam.; calyx lobes linear, ~1.2 mm long; corolla spinescens and Plumbago zeylanica. Undara Lava Tubes form rotate, (7–)10–13 mm in diam., 5-lobed, the lobes part of the Cainozoic McBride Volcanic Province (Griffin and rounded–triangular, 2–3.5 mm long; disc ~2 mm in diam.; McDougall 1975), the basalts of which provide habitat for other staminal column 1.8–2.2 mm long; staminal head 1.9–2.4 mm narrowly endemic plant species including Auranticarpa edentata in diam. Female inflorescence a (6–)9–14(–20)-flowered head; and Ipomoea saintronanensis. peduncle 12–35 mm long. Female flowers: subsessile; ovary ovate, 3.2–4 mm long, 2–2.6 in diam., echinulate with barbed Phenology aculei; hypanthium above the constriction broadly campanulate, Flowers and fruits March–April. ~2.4 mm in diam., abaxially scabridulous; calyx lobes linear, ~0.8 mm long; corolla 4–6.8 mm in diam., glabrous abaxially and Conservation status adaxially, glandular papillose on margins, white, 5-lobed; lobes – – Sicyos undara is known from two subpopulations at Undara, one triangular ovate, 2 2.3 mm long; disc ~1.3 mm in diam.; style with a single plant observed in March 2009, and the other with ~1.8 mm long; stigma 2-branched, the branches recurved. Fruit – – two. A survey is necessary to establish the range of the species and ovate, 8 13 mm long, 4 5.8 mm in diam., obtuse or subacute, size of populations. Until assessed, a threat category of ‘Critically the surface glabrous or sparsely and minutely hairy, echinate; – – Endangered’ is suggested under the Environment Protection aculei dense, (2.2 )3.6 6 mm long, retrorsely barbed. Seeds Æ – – and Biodiversity Conservation Act 1999 (Commonwealth). ellipsoidal, 5.6 6 mm long, 3.8 4.5 mm wide, brown. (Fig. 6). Too frequent wildfires would constitute a major threat. The species is conserved in Undara Volcanic National Park. Distribution Sicyos mawhai occurred formerly on mainland North Island Etymology and the Marlborough Sounds area of . Solander Named for the only known locality of the species, from the local (1782) recorded collections made in 1769–1770 on Captain aboriginal ‘a long way’. James Cook’s first voyage to the Pacific from North Island, including Tigadu (Anaura Bay), Tolaga (Tolaga Bay), Additional specimens examined Opuragai (Mercury Bay), Oohuragi (Firth of Thames, Waihou QUEENSLAND: Cook District: Yaramula Station [now Undara River), Motuaro (Bay of Islands, Motuarohia Island), and South Volcanic National Park], adjacent to Pinwill Cave, 80 km SW of Island, including Totaranui (Queen Charlotte Sound). The Mount Garnet, V.J. Neldner 2780 (BRI, CANB, CNS); Undara, Wind species is now confined to offshore islands, islets and rock New species of Sicyos in the South West Pacific Australian Systematic Botany 197

Fig. 6. Holotype of Sicyos mawhai I.Telford & P.Sebastian. stacks off New Zealand’s North Island from the Bay of Plenty forest (especially forest margins), coastal shrubland and on northwards, and on the more remote Three Kings and Kermadec offshore islands within sea-bird colonies, most notably, the Islands groups (Fig. 4). nesting grounds of petrels (Pterodroma spp., Puffinus spp.). Within its beach habitat, the species has been collected trailing Habitat and ecology through and over Coprosma repens, laetum, Ficinia Sicyos mawhai is a coastal species that has been collected from nodosa and Phormium tenax, and in forest it often associates beach strands, including boulder beaches and banks, from coastal with Corynocarpus laevigatus, Entelea arborescens, Myoporum 198 Australian Systematic Botany I. R. H. Telford et al. laetum, Melicytus ramiflorus, Macropiper excelsum subsp. Etymology excelsum, Pseudopanax lessonii and Metrosideros excelsa.On Named for the Maori name for the plant, mawhai, which means the Three Kings Islands, it often grows with Brachyglottis ‘wavy’ or ‘curly’, possibly in reference to the plant’s habit and/or aborescens, Coprosma repens, Metrosideros excelsa and tendrils. Mawhai is also said to mean ‘useless’ (de Lange 2011), Kunzea aff. ericoides (e), and at open sites with Ipomoea perhaps in reference to a Tainui legend. The name mawhai is also cairica. On the Kermadec Islands, it is a common species on widely applied to Cassytha paniculata R.Br. (de Lange 2011). Raoul Island where it mostly grows in canopy gaps and along forest–shrubland margins with Coprosma petiolata, Myrsine Illustrations kermadecensis, Myoporum rapense subsp. kermadecense, F.B. Sampson, Early New Zealand Botanical Art t. 28 (1985). Macropiper excelsum subsp. psittacorum, Metrosideros L. Forester & A. Townsend, Threatened Plants of Northland kermadecensis and Melicytus aff. ramiflorus. On the smaller Conservancy 61 (2004); two lower images. islands of the Kermadec group, it is also to be found trailing through petrel colonies, where it associates with Cyperus Additional specimens examined (selection) insularis, Scaevola gracilis, Solanum nodiflorum, Ipomoea cairica, I. pescaprae subsp. brasiliensis and Tetragonia NEW ZEALAND: Kermadec Islands: Raoul Island orchard, W.R. Sykes tetragonioides. Less commonly, like S. australis occurrences 188/K (CHR, K); Raoul Island, Fishing Rock, P.J. de Lange K490 (AK); Macauley Island, Mount Haszard, G. Taylor 13 (CHR); Macauley Island, in New Zealand proper, S. mawhai has been collected from fi Macauley Plateau, P.J. de Lange K817 (AK). Three Kings Islands: modi ed islands such as the Mokohinau Island group growing West Island, A.E. Wright 6061 (AK); Great Island, J.S. Edwards s.n. in induced pasture and wasteland, growing with grasses such as AK 44826 (AK). North Island: Te Paki, Kapowairua/Spirits Bay, Bromus willdenowii, Dactylis glomerata, bracken (Pteridium Maungapiko Hill, 29 September 2006, L. Bridson & W. Holland esculentum) and black berries (Rubus fruticosus agg.) (AK); Poor Knights Islands, Aorangi Island, Crater Bay, 2 January 1978, G.N. Park (CHR); Hen Island, February 1968, D. Merton Phenology (CHR); Little Barrier Island, near West Landing, W.R. Sykes 147/85 (CHR); Fanal Island, A.E. Wright 3146 (AK, HO, WELT); Coromandel Sicyos mawhai flowers and fruits mainly in December–May, and Islands, Motukarikitahi, A.E. Esler 3584 (CHR). CULTIVATED: Oratia also throughout the year in suitable conditions. Native Plant Nursery, Auckland, J.J. Bruhl 2915 (AK, CHR, M, NE).

Chromosome number Key to Sicyos in the South West Pacific region 2n = 26 (de Lange and Murray 2002). Male flowers mostly >10-mm diam...... S. mawhai Male flowers mostly <10-mm diam. Conservation status Fruit <6-mm long, attenuate; peduncle of female inflorescence mostly <25-mm long...... S. australis Cameron (1992) recorded the shrinking distribution of Fruit >6 mm long, apically rounded; peduncle of female inflorescence S. mawhai, with extinction at the southern end of its historical mostly >25 mm long...... S. undara range. No populations survive on mainland North or South Islands (P. J. de Lange, pers. comm.); note that the 1992 Acknowledgements Auckland collection cited by Cameron (1992) refers to We thank the directors of the herbaria AK, BM, BRI, CANB, CHR, HO, JCU, S. australis sensu stricto. Plants are susceptible to CMV and K, MEL, NSW and W for the loan of specimens and use of facilities, and the ZYMV (Delmiglio and Pearson 2006), and this appears to Queensland Department of Environment and Resource Management for weaken populations and has been used to explain the rapid permission to collect. We thank Werner Greuter for the impeccable Latin. decline of the species. Fruit and seedlings are eaten by Thanks go to John Hunnex of the Natural History Museum, London (BM), for rodents; removal of these vermin allows recolonisation an image of the Banks and Solander specimen. Ian R. H. Telford expresses (P. J. de Lange, pers. obs.). The species is rated ‘At Risk/ sincere thanks to Mike Crisp, Research School of Biology, The Australian Relict’ in de Lange et al.(2009b)asS. aff. australis (a) (AK National University, for his inspiration through time shared at the Australian 252822). National Botanic Gardens and as supervisor for the initial Sicyos morphometric analysis. Notes References Sicyos mawhai and S. australis occur almost sympatrically, but Allan HH (1961) ‘Flora of New Zealand.’ (Government Printer: Wellington, can readily be distinguished vegetatively; the former has leaves of New Zealand) a thicker texture, the lobes more rounded, with the basal pair Bailey FM (1900) ‘The Queensland flora.’ (Queensland Government: sometimes overlapping across the sinus, as well as by the floral Brisbane) and fruit differences presented in the key below. Beadle NCW (1976) ‘Students flora of north eastern New South Wales.’ Unfortunately, the Banks and Solander specimens housed in (Botany Department, University of New England: Armidale, NSW) ‘ the Museum of New Zealand Te Papa Tongarewa (WELT; http:// Beadle NCW, EvansOD, Carolin RC (1962) Handbookof the vascular plants of the Sydney District and the Blue Mountains.’ (Brown Gem Print: collections.tepapa.govt.nz/theme.aspx?irn = 2152, accessed 12 Armidale, NSW) October 2011) and the Natural History Museum, London Belbin L (1990a) ‘PATN: a user’s guide.’ (Division of Wildlife and Ecology, (BM), have no precise locality on the sheets. The single BM CSIRO: Canberra) specimen located has a quote from Solander (1782) attached Belbin L (1990b) ‘PATN: a technical reference.’ (Division of Wildlife and listing the collection sites cited above. Ecology, CSIRO: Canberra) New species of Sicyos in the South West Pacific Australian Systematic Botany 199

Belbin L, Collins A (2006) ‘PATN. V.3.1.’ (Blatant Fabrications Pty Ltd: Griffin TJ, McDougall I (1975) Geochronology of the Cainozoic McBride Brisbane) Volcanic Provinve, northern Queensland. Journal of the Geological Bentham G (1866) ‘Flora Australiensis.’ (Lovell Reeve & Co.: London) Society of Australia 22, 387–396. doi:10.1080/00167617508728905 Brandon A, de Lange P, Townsend A (2004) ‘Threatened plants of the HeenanPB,Forester LJ (1997)Mazus pumilio (),an addition Waikato Conservancy.’ (Department of Conservation: Wellington, to the indigenous flora of New Zealand. New Zealand Journal of Botany New Zealand) 35, 437–440. doi:10.1080/0028825X.1987.10410168 Buchanan AM (1988) The Tasmanian collecting localities of Ronald Gunn Heenan PB, de Lange PJ, Keeling JK (2009) Alternanthera nahui – a new and Joseph Milligan. Tasmanian Herbarium Occasional Publication 1, species of Amaranthaceae indigenous to New Zealand. New Zealand 1–56. Journal of Botany 47,97–105. doi:10.1080/00288250909509795 Cameron EK (1992) Decline of mawhai (Sicyos australis). New Zealand Hooker JD (1847) Flora Tasmaniae spicelegium. Hooker’s London Journal of Botanical Society Newsletter 28,11–12. Botany 6, 473–474. Chandler GT, Plunkett GM, Pinney SM, Cayzer LW, Gemmill CEC (2007) Hooker JD (1856) ‘The botany of the Antarctic voyages of H.M. discovery Molecular and morphological agreement in Pittosporaceae: phylogenetic ships Erebus and Terror. 3. Flora Tasmaniae.’ (Lovell Reeve & Co.: analysis with nuclear ITS and plastid trnL–tmF sequence data. Australian London) Systematic Botany 20, 390–401. doi:10.1071/SB07004 Hooker JD (1860) ‘The botany of the Antarctic voyage. 3. Flora Tasmaniae.’ Connor HE, Edgar E (1987) Name changes in the indigenous New Zealand (Lovell Reeve & Co.: London) Flora, 1960–1986, and nomina nova IV, 1983–1986. New Zealand Perrie LR, Brownsey PJ, Lockhart PJ, Brown EA, Large MF (2003) Journal of Botany 25, 115–171. doi:10.1080/0028825X.1987.10409961 Biogeography of temperate Australasian Polystichum ferns inferred Curtis WM (1963) ‘The Students flora of Tasmania.’ (Government Printer: from chloroplast sequence and AFLP. Journal of Biogeography 30, Hobart) 1729–1736. doi:10.1046/j.1365-2699.2003.00938.x de Lange PJ (1997) Gratiola pedunculata (Scrophulariaceae); a new addition Prebble JM, Cupido CN, Meudt HM, Garnock-Jones PJ (2011) First to the New Zealand flora. New Zealand Journal of Botany 35, 317–322. phylogenetic and biogeographical study of the southern bluebells doi:10.1080/0028825X.1997.10410157 (Wahlenbergia, Campanulaceae). Molecular Phylogenetics and de Lange PJ (2011) Cassytha paniculata a possible indigenous biocontrol for Evolution 59, 636–648. doi:10.1016/j.ympev.2011.03.013 gorse (Ulex europeus)? Trilepidea 93,9–10. Schaefer H, Renner SS (2011) Phylogenetic relationships in the order de Lange PJ, Murray BG (2002) Contributions to a chromosome atlas and a new classification of the gourd family of the New Zealand flora – 37. Miscellaneous families. New Zealand (Cucurbitaceae). Taxon 60, 122–138. Journal of Botany 40,1–23. doi:10.1080/0028825X.2002.9512767 Schaefer H, Heibl C, Renner SS (2009) Gourds afloat: a dated phylogeny de Lange PJ, Norton DA (1998) Revisiting rarity: a botanical perspective on reveals an Asian origin of the gourd family (Cucurbitaceae) and numerous the meanings of rarity and the classification of New Zealand’s uncommon oversea dispersal events. Proceedings. Biological Sciences 276, 843–851. plants. Royal Society of New Zealand Miscellaneous Series 48, 145–159. doi:10.1098/rspb.2008.1447 de Lange PJ, Norton DA, Heenan PB, Courtney SP, Molloy BPJ, Ogle CC, Sebastian P, Schaefer H, Telford IRH, Renner SS (2010a) Cucumber Rance BD, Johnson PN, Hitchmough R (2004) Threatened and (Cucumis sativus) and melon (C. melo) have numerous wild relatives uncommon plants of New Zealand. New Zealand Journal of Botany in Asia and Australia, and the sister species of melon is from Australia. 42,45–76. doi:10.1080/0028825X.2004.9512890 Proceedings of the National Academy of Sciences of the United States of de Lange PJ, Sawyer WD, Rolfe JR (2006) ‘New Zealand indigenous vascular America 107, 14 269–14 273. doi:10.1073/pnas.1005338107 plant checklist.’ (New Zealand Plant Conservation Network: Wellington, Sebastian PM, Schaefer H, Renner SS (2010b) Darwin’s Galapagos gourd: New Zealand) providing new insights 175 years after his visit. 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Freeman and Co.: San 61–96. doi:10.1080/00288250909509794 Francisco) de Lange PJ, Smissen RD, Wagstaff SJ, Keeling DJ, Murray BG, Toelken HR Solander DC (1782) Primitiae florae Novae Zelandiae. In ‘Floras of the (2010) A molecular phylogeny and infrageneric classification for Kunzea countries visited by Captain James Cook’s first voyage’. pp. 381–620. (Myrtaceae) inferred from rDNA ITS and ETS sequences. Australian (Botany Library, Natural History Museum: London) Systematic Botany 23, 309–319. doi:10.1071/SB10019 Stamatakis A (2006) RAxML-VI-HPC: maximum likelihood-based de Lange PJ, Townsend AJ, Rolfe JR (2011) Crassula nutans var. minus phylogenetic analysis with thousands of taxa and mixed models. (Crassulaceae) a new trans-Tasman natural weed arrival to northern Bioinformatics 22, 2688–2690. doi:10.1093/bioinformatics/btl446 New Zealand. New Zealand Journal of Botany 49, 361–366. 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Appendix 1. Dataset used in the phenetic analysis of Sicyos in Australia and the South West Pacific See Table 1 for voucher data. See Table 2 for characters relating to numbers in the header row. OTU, organisational taxonomic unit

OTU Character 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 NZ1 24 9 4.5 10.2 5.3 5.8 42 5.8 93 66 14 16.0 12.8 1.5 2.3 NZ2 23 8 4.2 10.0 5.4 5.4 50 6.0 135 95 25 10.9 8.7 1.5 2.4 NZ3 25 8 9.8 6.2 4.5 46 6.2 124 72 17 14.5 10.6 1.3 2.4 NZ4 26 8 4.3 8.5 6.0 4.2 53 6.4 152 85 35 11.0 10.4 1.8 2.2 NZ5 25 8 4.8 9.2 6.2 5.3 48 5.5 110 72 18 12.4 9.2 1.5 2.0 NZ6 14 10 7.8 3.8 4.8 26 4.8 37 22 8 4.8 NZ7 12 7 3.0 7.2 3.2 4.2 27 4.2 53 42 6 5.0 NZ8 25 9 10.5 6.5 4.6 52 6.5 140 88 19 12.0 10.0 2.2 2.8 NZ9 12 10 4.3 8.0 3.8 4.2 20 4.0 55 22 11 4.2 8.2 1.4 1.7 NZ10 16 8 8.8 3.6 3.8 17 4.8 106 68 21 4.5 NZ11 24 14 4.0 11.3 6.5 4.5 38 6.5 116 65 22 11.2 11.8 NZ12 34 15 4.8 11.8 6.7 5.5 27 104 78 15 12.5 10.0 KI1 24 8 4.5 11.8 6.0 5.5 36 6.8 110 79 16 12.3 10.5 KI2 18 8 9.0 5.8 4.6 40 6.0 92 84 23 9.5 10.6 1.3 2.4 KI3 23 9 9.2 5.2 4.8 56 6.5 126 52 22 10.8 10.2 2.0 1.9 NI1 23 9 2.6 7.8 3.7 7.0 25 4.8 124 93 13 9.3 4.5 NI2 16 8 2.5 7.2 3.2 5.5 24 61 49 9 6.5 5.0 LH1 22 10 3.0 7.8 3.5 6.5 24 4.7 62 47 8 6.2 6.8 LH2 28 8 7.2 3.3 5.5 28 78 61 10 6.0 6.2 LH3 17 7 7.0 3.0 6.9 26 4.4 80 36 14 5.8 5.8 TS1 10 8 3.4 7.8 3.7 6.0 23 57 35 11 5.2 6.3 1.8 2.2 TS2 11 10 2.8 7.3 3.5 7.5 22 4.6 68 40 12 3.5 6.0 VG1 21 7 2.7 7.2 3.5 6.0 23 4.5 79 67 12 5.0 6.5 VG2 14 8 2.5 6.5 3.0 5.8 21 67 50 15 3.5 5.8 1.5 1.8 NS1 13 6 5.5 2.7 4.5 19 4.0 60 53 7 4.8 7.4 1.8 1.7 NS2 23 7 2.8 7.5 3.6 7.0 28 4.8 104 93 12 8.5 6.4 1.5 1.7 NS3 15 8 2.4 8.2 3.8 3.5 22 5.0 87 65 11 6.8 6.0 QL1 13 10 2.4 7.0 3.7 3.5 28 4.2 90 78 13 6.5 5.3 QL2 15 6 3.0 7.5 3.8 6.5 21 79 55 9 4.5 5.8 QL3 17 7 6.5 3.5 6.8 24 4.0 114 89 11 6.8 5.3 QL4 11 7 2.5 8.3 4.0 7.0 22 93 75 12 4.0 4.2 1.6 1.7 QL5 36 5 2.0 4.8 2.8 2.6 13 3.5 60 37 14 9.6 4.2 1.1 0.8 QL6 27 4 5.3 3.0 2.8 11 34 27 12 6.0 3.7 0.9 0.7 QL7 57 6 2.4 4.7 2.9 2.8 12 3.2 95 22 11.3 4.8 0.9 0.7

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