Research Article |
Corresponding author: Peter de Lange ( pj.delange@xtra.co.nz ) Academic editor: Marcin Nobis
© 2016 Peter de Lange, Rob D. Smissen, Jeremy R. Rolfe, Colin C. Ogle.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
de Lange PJ, Smissen RD, Rolfe JR, Ogle CC (2016) Systematics of Simplicia Kirk (Poaceae, Agrostidinae) – an endemic, threatened New Zealand grass genus. PhytoKeys 75: 119-144. https://doi.org/10.3897/phytokeys.75.10328
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A new species of the New Zealand endemic grass Simplicia, S. felix is described. The new species is segregated from and compared with S. buchananii and S. laxa. Simplicia felix occurs mostly in lightly shaded areas of seasonally dry alluvial forest. A distribution map and an assessment of the conservation status of the new species are presented. Genetic variation in the genus was examined, building on previously published work but including additional sampling. Analysis of nrDNA ITS and ETS and plastid trnL intron and trnL–F intergenic spacer sequences show S. felix to be more closely related to S. laxa than to S. buchananii. NeighborNet analyses of AFLP profiles for the three species of Simplicia show each to consist of distinct clusters of genotypes well separated from each other.
Poaceae , Simplicia , new species, S. felix , S. buchananii , S. laxa , conservation status, New Zealand flora
During 2005 one of us (Colin Ogle), chanced upon an unremarkable grass growing in an alluvial forest remnant on the margin of a grazed pasture in the Kawhatau Valley Mangaweka, in the Rangitikei District, North Island (
To resolve the matter a DNA-based investigation of the genus was initiated to try to determine which species the plant actually was (
During February 2014 Simplicia was rediscovered in the Wairarapa (
That data, along with morphological evidence, suggests that the recognition of a third species of Simplicia and a narrower circumscription of S. laxa are justified.
Details of samples and GenBank accessions are shown in Table
Species | Location | Region | Lat/long | vouchers | GenBank ITS |
GenBank ETS |
GenBank trnL/F |
---|---|---|---|---|---|---|---|
Simplicia buchananii | Cobb Valley | Nelson | 41°7'50.96"S, 172°36'31.53"E | No Voucher | HM191441 | HM191453 | HM191463 |
Simplicia buchananii | West of Gordon’s Pyramid | Nelson | 41°11'15.24"S, 172°40'3.32"E |
|
HM191439 | HM191451 | HM191465 |
Simplicia buchananii | Cundy Creek | Nelson | 41°11'14.32"S, 172°40'02.28"E |
|
HM191440 | HM191452 | HM191464 |
Simplicia felix | Williams’s Bush | Wellington | 39°37’ 20.7"S, 175°45'54.02” E |
|
HM191437 | HM191449 | HM191461 |
Simplicia felix | Taihape Reserve | Wellington | 39°40'14.29"S, 175°48'18.96"E |
|
HM191435 | HM191447 | HM191459 |
Simplicia felix | Ben Moi Farm | Wellington | 39°46'50.25"S, 175°51'59.14"E |
|
HM191438 | HM191450 | HM191462 |
Simplicia felix | Kaumingi 1 | Wellington | 40°57’ 50.18"S, 175°52’ 15.83"E |
|
KU724189 | KU728132 | KU728136 |
Simplicia felix | Kaumingi 2 | Wellington | 40°58’ 4.56"S, 175°52’ 26.49"E |
|
KU724190 | KU728131 | KU728135 |
Simplicia felix | Te Kowhai | Wellington | 41°10’ 57.62"S, 175°40’ 59.85"E |
|
KU724191 | KU728130 | KU728134 |
Simplicia laxa | Honeycomb Cave | Nelson | 41°8'25.79"S, 172°11'42.41"E |
|
HM191442 | HM191454 | HM191466 |
Simplicia felix | Ngapara | Otago | AK285424 | KU724188 | KU728129 | KU728133 | |
Simplicia laxa | Nenthorn | Otago | 45°28'39.6"S, 170°22'47.3"E |
|
HM191445 | HM191457 | HM191470 |
Simplicia laxa | Macraes | Otago | 45°25'35.95"S, 170°27'13.9"E |
|
- | - | - |
Simplicia laxa | Castle Rock | Otago | 45°17'53.92"S, 169°16'13.87E |
|
HM191443
HM191446 |
HM191455
HM191458 |
HM191467
HM191469 |
For DNA sequencing methods see
New AFLP profiles were generated from the same three Wairarapa samples plus an additional sample (see Table
Descriptions of S. buchananii, S. laxa sens. str. and the new species were prepared using cultivated plants and herbarium specimens held at
Little resolution was obtained by phylogenetic analysis of the plastid DNA sequences which included only 12 variable characters (four with outgroups excluded), only five of which were parsimony informative (three with outgroups excluded). In the single most parsimonious tree (length 13) found by heuristic search of these (not shown) the three S. buchananii samples are characterised by two synapomorphic substitutions in the region sequenced. Some but not all of the North Island S. laxa samples share a single synapomorphic substitution.
More resolution was obtained in analysis of the nrDNA sequences which contained 196 variable characters (25 with outgroups excluded), 53 of which were parsimony informative (20 with outgroups excluded). Three equally parsimonious trees (length 218) were recovered by the heuristic search. One of these (Fig.
Fragment scoring with the settings described above resulted in data matrices with 475 fragments for Eco-AAC with M-CTC and 559 for Eco-ACG with Mse-CAG. NeighborNet graphs generated for each primer pair showed similar grouping of samples (not shown) and we combined them to form a single data matrix with 1034 fragments. The NeighborNet for the full data matrix is shown in Fig.
Three major groups are obtained in both NeighborNet analyses of AFLP profiles. One group corresponds with S. buchananii including samples from Gordons Pyramid, Cobb Valley, and Cundy Creek (see Fig.
Relative to eyeball scoring of silver-stained slab gels, fluorescent labelling and automated scoring AFLP profiles produces many more apparent polymorphisms but with considerably reduced profile reproducibility, at least for Simplicia in our hands (see
The results reported here are in accord with those obtained by
Seed set in isolated, glasshouse-grown plants indicate that both S. buchananii and S. laxa are self-compatible (
S. laxa Kirk (fide
1 | Plants tufted; culms erect, up to 1 m tall (culm nodes not root-forming); inflorescences erect, linear, branches, erect, appressed to rachis, bearing spikelets almost to base | S. buchananii |
– | Plants decumbent; culms weakly ascendant, rooting freely from culm-nodes so forming diffuse interconnected widely sprawling clonal patches 0.6–1.0 m diameter; inflorescences linear to pyramidal, binate, basal branch or branches reflexed, devoid of spikelets from lower ½ to ⅔ | 3 |
2 | Mid-stem and upper stem leaf sheaths finely ribbed, copiously hairy (hairs 0.35–0.40 mm long); adaxial leaf-blade ribs hairy; inflorescence branches antrorsely hairy, pedicels 1.00–1.06 mm long; lemma pubescent | S. laxa |
– | Mid-stem and upper stem leaf sheaths strongly ribbed, ± glabrous (occasionally bearing minute hairs towards sheath apex); adaxial leaf-blade ribs smooth or finely scabrid; inflorescence branches scabrid, pedicels 0.20–0.30 mm long; lemma minutely scabrid | S. felix |
≡ Poa uniflora Buchanan Indig. Grasses N.Z. t49B (1880) non Muhl. (1817)
≡ Simplicia laxa var. buchananii Zotov T.R.S.N.Z. 73: 236 (1943)
‘Mt Arthur’ A. McKay s.n., 1874, (
Named by
(Fig.
North-West Nelson, Gouland Downs, A.P. Druce s.n., Jan 1969,
(Fig.
Simplicia buchananii is distinguished from S. laxa and S. felix by the tufted growth habit, erect culms, and by the linear inflorescences, whose branches are usually tightly appressed to the rachises (Fig.
Simplicia buchananii is a biologically sparse species of calcareous rock habitats (including shaded rock outcrops, boulderfalls, rock overhangs and cave entrances) within montane forest (
Using the New Zealand Threat Classification System (
≡ Simplicia laxa var. laxa (autonym, Zotov T.R.S.N.Z. 73: 236 (1943))
‘Waikouaiti, Otago’ D. Petrie s.n., n.d. (
‘Waikouaiti, Otago, D.Petrie s.n., n.d. (
(Fig.
Simplicia laxa. A habitat, Emerald Stream, McCraes, North Otago, South Island, New Zealand (image D.A. Houston) B growth habit and inflorescence C culm, leaf base, sheath and ligule D spikelet showing reduced glumes and lemma. (Photo credit images C and D: K. Ford, Allan Herbarium, Landcare Research Manaaki Whenua)
Karamea, Honeycomb Cave, P. Wardle s.n., 22 Jan 1985,
(Fig.
Distinguishing features of Simplicia species based on wild collected material.
Simplicia buchananii | Simplicia laxa | Simplicia felix | |
---|---|---|---|
Growth habit | Tufted, erect. culms up to 1 m tall | Decumbent, culms sprawling, forming mats up to 0.6 m diameter | Decumbent, sprawling, forming mats up to 1 m diameter |
Culm internodes | Glabrous, ± equal in length to subtending leaf sheaths | Hairy or glabrous, < subtending leaf-sheaths. Hairs if present up to 0.18 mm long | Glabrous, > subtending leaf-sheaths |
Culm nodes | Glossy orange-brown to dark red-brown | Glossy maroon-black to black | Glossy dark brown-green to brown-black |
Basal leaf sheaths | Strongly ribbed, stramineous or dull brown, glabrous or with ribs scabrid (very rarely finely pubescent). Hairs (if present) 0.06–0.08 mm long | Finely (‘weakly’) ribbed, glossy light brown to amber, ribs pubescent. Hairs 0.20–0.30 mm long | Strongly ribbed, dull dark brown, ribs glabrous or pubescent. Hairs (if present) 0.10–0.15 mm long |
Mid-stem and upper-stem leaf sheaths | Strongly ribbed. Glabrous | Hairy. Hairs copious, 0.35–0.40 mm long | Strongly ribbed. Usually glabrous, occasionally ribs finely short pubescent towards sheath apex |
Ligule | Glabrous | Sparsely to copiously hairy. Hairs 0.20–0.24 mm long | Glabrous or with both surfaces finely hairy. Hairs 0.15–0.18 mm long |
Leaf-blade | 1.8–4.0 mm wide. Adaxially finely scabrid, abaxially glabrous, margins smooth. Apex acuminate | 2.8–3.6 mm wide. Adaxially with hairy ribs, abaxially glabrous; margins ± smooth, sometimes, finely scabrid and/or sparsely hairy. Apex acute | 1.0–3.0 mm wide. Ribs (both surfaces) smooth or finely scabrid; margins smooth or finely scabrid. Apex acute |
Inflorescence | Paniculate, linear up to 180 mm long. Branches glabrous, erect, appressed to rachis, bearing spikelets almost to base. | Paniculate, linear to ± pyramidal, up to 150 mm long. Branches antrorsely hairy, basal branches (or branch) reflexed, others weakly appressed to rachis, lower half of branch devoid of spikelets | Paniculate, linear to ± pyramidal, up to 80 mm long. Branches scabrid, basal branches (or branch) reflexed, others weakly appressed to rachis, lower ½ to ⅔ or branch devoid of spikelets |
Pedicels | Glabrous, 0.60–1.2 mm long | Pubescent, 1.00–1.06 mm long | Pubescent 0.20–0.30 mm long |
Glumes | Lower glume 0.3–1.0 mm long, upper glume 1.0–1.6 mm long | Lower glume 0.5–0.8 mm long, upper glume 0.75–1.2 mm long | Lower glume 0.5–0.6 mm long, upper glume 0.75–0.9 mm long |
Lemma | 2.6–3.0 mm long, scabrid | 2.8–3.4 mm long, pubescent | 2.0–3.0 mm long, minutely scabrid |
Rhacilla prolongation | 0.3–0.5 mm long, filiform, glabrous | 1.25–1.30 mm long, narrowly lanceolate, margins minutely ciliate | 0.8 mm long, filiform, glabrous except for sparse cilia cresting apex |
Anther filaments | 0.3 mm long, | 0.20–0.25 mm long | 0.6–0.9 mm long |
Anthers | 0.7–1.5 mm long, | 0.30–0.45 mm long | 1.0–1.2 mm long |
Because of the lax, decumbent trailing growth habit, and loose linear to pyramidal inflorescences Simplicia laxa is easily distinguished in the field from the shortly tufted S. buchananii whose inflorescences are erect and whose inflorescence branches are held tightly appressed to the rhacis (
Much of what has been written about Simplicia laxa (
Simplicia laxa was assessed as ‘Threatened / Nationally Critical’ qualified ‘CD’ (Conservation Dependent), ‘Sp’ (Sparse) by de Lange et al. (2013). That assessment included plants described here as Simplicia felix. With the recircumscription of S. laxa the species remains appropriate assessed as ‘Nationally Critical’. Currently there are < 15 populations known, and several of these are in decline, and very few are substantial in size. Many occur on private land without direct conservation management or in places subject to ongoing habitat deterioration through invasive weed pressure and habitat loss. The qualifiers however need adjustment. Because the species is managed in a number of sites it is appropriate to retain the qualifier ‘CD’, as ceasing management would have a serious impact on the survival of the species. It is debatable whether Simplicia laxa is truly biologically sparse. It is sparsely distributed but this is more likely an artefact of past habitat loss leaving highly fragmented, disjunct ‘remnant’ populations rather than any natural pattern of distribution or species biology. We recommend that ‘Sp’ be removed from the conservation assessment for this species. The seemingly peculiar North-West Nelson, Honeycomb Cave outlier suggests that Simplicia laxa should be looked for throughout the South Island rather than, as it currently has, only in the Central and Eastern Otago Region. Also, it is now evident that we lack trend data for the species, though the overall impression is that many populations are in decline. For these reasons, we recommend that the species be qualified ‘DP’ (Data Poor) be added to the species conservation status.
Differs from Simplicia laxa by dark brown, prominently ribbed leaf sheaths; mostly glabrous, strongly ribbed mid-stem to upper-stem leaf sheaths; longer culm internodes; narrower, glabrous (sometimes with the adaxial ribs finely scabrid) leaves; shorter panicles (up to 80 mm long) with scabrid branches; minutely scabrid lemma and smaller filiform rachilla prolongation bearing cilia only at the apex.
The epithet ‘felix’ is taken from the Latin for ‘lucky’ (N.G. Walsh, MEL pers comm., 14 January 2016) as in ‘lucky find’ in reference to the circumstances of this species’ discovery; that came about through the desire to get a name on an unremarkable little tuft of grass that was discovered fortuitously near Mangaweka, Central North Island by CCO on 29 January 2005 (
(Fig.
New Zealand, North Island: Rangitikei Ecological Region and District, North of Taihape, north of Paengaroa Road and east of State Highway One, Ngawaka Stream, ‘Stevies Bush’, P.J. de Lange 7834 & C.C.Ogle, 29 Feb 2008,
(Fig.
Genetically and morphologically Simplicia felix is more closely related to S. laxa than it is to S. buchananii. From Simplicia laxa it can be distinguished by the culm internodes which are longer than the subtending leaf sheaths and consistently glabrous; and by the strongly ribbed, dull dark brown basal sheaths, whose ribs are glabrous or pubescent (if pubescent then with the hairs 0.10–0.15 mm long). The mid-stem and upper-stem leaf sheaths of S. felix are strongly ribbed and usually glabrous (occasionally the ribs are finely pubescent towards the sheath apex). The leaf surfaces and margins of S. felix are mostly smooth though the ribs and leaf margins may be minutely scabrid. As a rule, the leaves of S. felix are also shorter (up to 60 mm in S. felix, 200 mm in S. laxa) and narrower than those of S. laxa (1.0–3.0 mm long cf. 2.8–3.6 mm long in S. laxa). However, in cultivation the leaves of both species can get up to 4.0 mm wide. Although the inflorescences of S. laxa and S. felix are similar, those of S. felix are smaller (up to 80 mm long rather than 150 mm long), and the branches are scabrid rather than antrorsely hairy. Although the lemma of both species overlap in range, those of S. felix tend to be shorter (2.0–3.0 mm long) than those of S. laxa (2.8–3.4 mm long) and minutely scabrid rather than pubescent. The rachilla prolongation of S. laxa is narrowly lanceolate, 1.25–1.30 mm long and with the margins minutely ciliate, while that of S. felix is filiform, 0.8 mm long and bearing sparse cilia only near the apex. Other differences are given in Table
Simplicia felix was initially confused with S. buchananii (see
With the exception of the Ngapara population which grows within a limestone overhang, Simplicia felix has only been collected from central and eastern North Island lowland to lower montane, often riparian, seasonally dry (drought prone), Podocarp forests overlying base-rich substrates such as limestone, calcareous mudstone and siltstone. In these areas plants have been found only in lightly shaded situations within forest remnants that have been lightly under-grazed by cattle or sheep (
The Ngapara site, as S. laxa, was described in some detail by
Simplicia felix occupies a very small area of only a few square metres wherever it occurs. It appears to have quite specific light requirements and tolerates only limited competition from other ground-cover species. The healthiest populations occur at sites where competition is reduced by grazing mammals such as cattle and sheep. This poses a quandary for conservation managers because, whilst grazing apparently benefits S. felix, it will ultimately lead to the collapse of the forest canopy that provides the level of shade that is also necessary for S. felix to survive. Collectively, the area of the sites where S. felix occurs amounts to considerably less than 1 ha. Therefore, S. felix meets the criteria to be assessed Threatened—Nationally Critical B2 using the New Zealand Threat Classification System (
The authors would like to thank Dave Houston, Mike Thorsen, Peter Johnson, Tony Silbery, the late Phil Knightbridge, Graeme Wood, John Barkla, Rhys Gardner, Kerry Ford, Robert Soreng, Neville Walsh, Geoff Davidson, Viv McGlynn, Graeme La Cock and Phyllis Leigh for company in the field and sharing their knowledge of Simplicia. We thank Ewen Cameron (