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The genus Empodisma comprises two species that are ecologically important in wetland habitats. Empodisma gracillimum is restricted to south-western Australia, whereas Empodisma minus is found in Tasmania, eastern Australia and New Zealand. We sequenced three cpDNA genes for 15 individuals of Empodisma sampled from throughout the range of the species. The results support an Australian origin for Empodisma sometime during the late Oligocene to early Miocene with more recent dispersal, colonization and diversification in New Zealand. We recovered six genetically distinct maternal lineages: three Empodisma gracillimum haplotypes corresponding to the three accessions in our analysis, a wide-ranging Empodisma minus haplotype found in eastern Australia and Tasmania, an Empodisma minus haplotype found in New Zealand from Stewart Island to approximately 38° S latitude on the North Island, and a distinct haplotype restricted to the North Island of New Zealand north of 38° S latitude. The Eastern Australian and New Zealand haplotypes of Empodisma minus were supported by only one cpDNA gene, and we felt the relatively minor morphological differences and the small amount of genetic divergence did not warrant taxonomic recognition. However, we recommend that the northern New Zealand haplotype should be recognized as the new species Empodisma robustum and provide descriptions and a key to the species of Empodisma. Monophyly of Empodisma robustum is supported by all three cpDNA genes. Empodisma robustum can be distinguished from Empodisma gracillimum and Empodisma minus by its robust growth stature and distinct ecology. It is typically eliminated by fire and re-establishes by seed (seeder strategy), whereas Empodisma minus and Empodisma gracillimum regrow after fire (sprouter strategy).
Restionaceae, Empodisma, taxonomy, new species, New Zealand
As presently circumscribed, the genus Empodisma L.A.S.Johnson & D.F.Cutler (Restionaceae) comprises two species with a widely disjunct distribution in western Australia and south eastern Australia, Tasmania, and New Zealand. Empodisma gracillimum (F.Muell.) L.A.S.Johnson & D.F.Cutler is found on the coastal plain from Perth southwards, and along the south coast from Augusta to Albany (Fig. 1), while Empodisma minus (Hook.f.) L.A.S.Johnson & D.F.Cutler is found in lowland to alpine zones from Queensland to South Australia, Tasmania and throughout most of New Zealand in New Zealand. They probably diversified in seasonally wet habitats, but exhibit adaptations to seasonal drought, fire and nutrient poor soils (
Map showing the generalized distribution of Empodisma in Australia and New Zealand and the collection localities of the DNA samples included in our study. The approximate position of the kauri line in New Zealand is shown with a dashed line.
The species of Empodisma are plants of peatlands, particularly raised bogs, blanket bogs, fens, and wet heathlands (
Fire plays an important role in the development of restiad peat bogs in both Australia and New Zealand. For the most part, the species of Empodisma are “sprouters” (
The taxonomic history of Empodisma is complex. The species of Empodisma were originally placed in Calorophus Labill or Hypolaena R.Br. by early taxonomists classifying Restionaceae (
The specimen upon which Hooker based the name Calorophus minor is a small slender plant characteristic of alpine regions found in South Island and Stewart Island of New Zealand.
We conducted a global analysis of 48 members of the Restionaceae to test monophyly of Empodisma and its relationships to Calorophus and Hypoleana. Three genera of Anarthriaceae (Anarthria R. Br., Hopkinsia W. Fitzg. and Lyginia R. Br.) were selected as outgroups. Intraspecific variation within Empodisma was assessed by comparing DNA sequences from 18 accessions collected from throughout the range of these species (Fig. 1).
Data resourcesVoucher specimens with their collection locality and GenBank accession numbers are listed in Appendix 1. The aligned data matrices have been submitted in Nexus format to TreeBase matrix accession number http://purl.org/phylo/treebase/phylows/study/TB2:S12748 and the Dryad repository: https://doi.org/10.5061/dryad.94710.
Morphological analysesA single set of morphological measurements were taken from each of 76 dried herbarium specimens on loan from AK, CHR, WAIK, WELT and PERTH (abbreviations follow Index Herbariorum). The morphological measurements describe the growth habit and floral structures of Empodisma and were confirmed with additional observations from field collections. Because Empodisma is dioecious, floral attributes of female flowers were coded as missing on male plants. Also very few specimens had mature fruits, and this attribute was also coded as missing from many specimens. We used GenStat version 8.1.0.152 (supplied by VSN International Ltd., www.vsn-int.com) to illustrate patterns of variation among the characters (listed in Fig. 6) using BOXPLOTS and PRINCIPAL COORDINATES ANALYSIS (PCoA). Principal coordinates analysis depicts relationships among the 91 Operational Taxonomic Units (OTU’s) that comprised our sample. We initially generated a similarity matrix of Euclidean distances then created a two-dimensional ordination. The first axis accounted most of the variation with less variation described by the second axis.
To test the influence of environmental conditions on growth form, we set up a common garden experiment by transplanting individuals (n=3) from New Zealand sites representing populations of both large (Empodisma robustum)and small (Empodisma minus)growth forms to Hamilton, North Island (37°47'S latitude). The sites selected were Torehape (37°18'S), Kopuatai (37°24'S), and Moanatuatua (37°55'S) for the large growth form, and Tongariro (39°16'S), Rangipo (39°22'S), and Awarua (46°33'S) for the small form. Ecological information was summarized from the published literature and unpublished data of BRC.
DNA extraction, amplification and sequencingWe extracted total DNA from either freshly collected plants or plants dried using silica gel, using a Qiagen DNeasy extraction kit (QIAGEN Pty Inc., Clifton Hill, Victoria, Australia) following the manufacturer’s directions. Three chloroplast-encoded DNA regions were sequenced: rbcL, matK and trnL. These regions were selected as they have been used previously to resolve relationships within the Restionaceae (
Our PCR amplification and sequencing procedures generally followed those described by
We used ClustalX (
Closely related sequences were initially aligned using the multiple alignment settings, a gap opening penalty of 5, a gap extension penalty of 5, and a delay-divergent-sequences setting of 97%. These ClustalX penalties favour opening gaps rather than substitutions, and they delayed adding the most distantly related taxa in our study. We identified low-scoring segments and exceptional residues, using the quality settings in ClustalX, and reconciled alternative alignments of these short DNA stretches. The final alignments were then visually inspected, and minor adjustments were made manually before conducting the phylogenetic analyses.
Some of the outgroup sequences were not available from GenBank (e.g. a matK sequence was missing for Chordifex hookeri (D.I.Morris) B.G.Briggs, but both an rbcL and trnL sequence were available from GenBank for this taxon). Rather than excluding these taxa, the incomplete data partitions were coded as missing. Many recent empirical and simulated studies suggest that it is possible to include taxa with large amounts of missing data without compromising phylogenetic accuracy. Indeed, increasing both the number of taxa and characters can improve the accuracy of phylogenetic inferences (
We conducted both parsimony and network analyses of the sequence data sets, using PAUP* 4.0b10 (
Each gene partition was tested for the best substitution model using jModelTest (
The earliest verifiable fossils of Poales are from the early Cretaceous (Maastrichtian) deposits dated approximately 115 million years ago (
The combined sequence data set comprised three data partitions with a total of 4267 characters; approximately 23% of the total data matrix was comprised of gap or missing data. An ILD test of the three data partitions failed to find significant conflict (p = 1–82/100 = 0.18). A heuristic search with parsimony as the optimality criterion recovered a single island of 180 trees of 2226 steps (Consistency Index (CI) = 0.607 (excluding uninformative characters); Retention Index = 0.788); a strict consensus is shown in Fig. 2. The three gene regions differed in length, the number of variable characters, and the degree to which they resolved and supported phylogenetic relationships.
Strict consensus tree. The three species of Empodisma (highlighted in bold) emerge as a well-supported clade distinct from Calorophus and Hypolaena. They were placed in these latter two genera by
The strict consensus tree (Fig. 2) agrees with the subfamilial classification of
The maximum parsimony and Bayesian analyses converged on trees with essentially the same topology, which suggested that the sequence data are robust to the different assumptions associated with these two approaches. Notably, the Bayesian posterior probability values were generally higher than the bootstrap support values, and the chronogram was better resolved (Fig. 3). The 95% highest posterior density estimates revealed substantial uncertainty associated with the divergence estimates, so our results should be viewed as preliminary. The results suggest Empodisma diverged from its most closely related ancestor (MCRA), Winifredia sola L.A.S.Johnson & B.G.Briggs approximately 21.8 (15.9–28.2) million years ago (mya). Empodisma gracillimum diverged at about 8.8 (5.4–12.9) mya and Empodisma robustum split from Empodisma minus approximately 2.0 (0.8–3.8) mya. Even though the algorithm for dating divergence times was different, the estimates presented here are similar to those obtained by
Bayesian chronogram with estimated divergence times. Node error bars are provided in blue showing the 95% highest probability density for the divergence estimates. Posterior probability support values > 97% are given above the branches. A geological time scale is shown at the base of the tree.
A comparison of median networks assessing levels of intraspecific variation in the three species of Empodisma is shown in Fig. 4. In each instance only a single parsimony tree was recovered. The trnL sequences were the shortest but the most variable. They were 953 nucleotides in length, and of these, 18 substitutions were parsimony informative. Fourteen substitutions supported the Empodisma gracillimum lineage with two unique parsimony uninformative substitutions distinguishing Empodisma gracillimum9.14 from the other species. One trnL character supported the eight accessions of Empodisma minus (bootstrap 63%). The split between the Australian (Empodisma minus7.45 and Empodisma minus9.15) and the New Zealand specimens (Empodisma minus8.09, Empodisma minus9.05), was supported by one trnL character but again with low support (65% bootstrap). Two informative substitutions supported the split between the robust northern New Zealand specimens of Empodisma (e.g. Empodisma robustum9.06 and Empodisma robustum7.44) highlighted in bold and the other specimens of Empodisma in our data set (Fig. 4). Further support for this split comes from a 23-base duplication that is absent from Empodisma robustum but present in Empodisma minus and Empodisma gracillimum. This split received 86% bootstrap support in our analysis. The sequences were identical within the seven accessions of Empodisma robustum.
Comparison of median networks from independent analyses of trnL, rbcL and matK sequences. Bootstrap values / the number of mutations distinguishing each haplotype. are shown beside the branches. The accessions of Empodisma minus from New Zealand are indicated NZ and Australia Aus.
The rbcL sequences were 1401 nucleotides long; of these, eight characters were parsimony informative and 1393 were constant. The informative characters support four splits in the data (Fig. 4). Five substitutions support Empodisma gracillimum with one substitution supporting Empodisma gracillimum12.1 and Empodisma gracillimum12.2. The third split supports only the diminutive accessions of Empodisma minus from New Zealand (65% bootstrap), and one substitution supports the fourth split separating the large lowland form of Empodisma (e.g. Empodisma robustum7.43 highlighted in bold (68% bootstrap).
By comparison, the matK sequences were 1469 nucleotides long, and of these 16 characters were parsimony informative, 3 variable characters were parsimony uninformative, and 1450 were constant. The informative characters again provided strong support for the split between Empodisma gracillimum and the remaining samples in our data (14 substitutions/100% bootstrap) (Fig. 4). Empodisma gracillimum12.1 was supported by two unique substitutions and Empodisma gracillimum9.14 by one. The split between specimens of Empodisma robustum from northern New Zealand and Empodisma minus was supported by 2 substitutions / 86% bootstrap. The sequences within these latter two groups were identical.
The Incongruence Length Difference test failed to reveal significant incongruence (p = 1–1/100 = 1.00) among the three independent data sets, so we pooled them. An analysis of the combined data recovered a single maximum parsimony tree; an unrooted phylogram is shown in Fig. 5 (Consistency Index, excluding uninformative characters = 1.00, Retention Index = 1.00). The combined analysis provided strong support for clades corresponding to the western Australian endemic, Empodisma gracillimum (33 substitutions / 100% bootstrap) and the robust northern New Zealand plants (e.g. Empodisma robustum9.06 and Empodisma robustum9.01 (five substitutions / 100% bootstrap), but weak support for Empodisma minus (1 substitution / 63% bootstrap).
Unrooted parsimony tree from a combined analysis of the three sequence partitions. Six distinct cpDNA haplotypes are supported. Bootstrap values / the number of mutations distinguishing each haplotype are shown beside the branches. The accessions of Empodisma minus from New Zealand are indicated NZ and Australia Aus.
We distinguished six distinct cpDNA haplotypes within the three species of Empodisma. Each accession of Empodisma gracillimum was distinguished by one or more substitutions and constituted three unique haplotypes. The two sequences of Empodisma minus from eastern Australia and Tasmania were identical and comprised the fourth haplotype (see also Figure 1). These were distinguished from a fifth haplotype comprising the Empodisma minus accessions from New Zealand ranging from Stewart Island to approximately 38° S latitude on the North Island. The sequences of Empodisma robustum restricted to the North Island of New Zealand north of 38° S latitude comprised a sixth haplotype.
We observed also a substantial degree phenotypic variation within the species of Empodisma especially in those characters that describe growth habit, e.g. culm height, internode distance, sheath length and leaf length. However, when grown together in common garden experiments in Hamilton, the two New Zealand species, Empodisma robustum and Empodisma minus, retained their distinctive growth forms, which suggests there is a genetic component to the pattern of morphological variation.
Empodisma robustum is generally a larger more robust plant, which ranges in height from 0.4 to over 1.3 meven taller in supporting vegetation, whereas Empodisma minus approaches 0.8 m in lowland bogs in Queensland, but in southern latitudes and alpine environments the plants are dwarfed, barely reaching 0.3 m (Fig. 6). Empodisma gracillimum is similar in height to Empodisma robustum, but the culms are light green in colour and more delicate; usually they are less than 0.7 mm in diameter. The culms of Empodisma robustum are dark green and broader, in some individuals approaching 2.2 mm in diameter. The culms of Empodisma minus are also dark green, but they are seldom greater than 1.0 mm in diameter. Internode distances also vary substantially among the three species; the distances are greater in Empodisma robustum and Empodisma gracillimum ranging from 20.0 to 70.0 mm in Empodisma robustum and from 25.0 to 80.0 in Empodisma gracillimum in contrast to Empodisma minus which ranges from 15.0 to 48.0 mm (Fig. 6). The leaf sheaths of Empodisma robustum also tend to be longer, ranging from 5.2 to 21.0 mm, whereas the leaf sheaths of Empodisma gracillimum range from 3.5 to 9.3 mm in length andfrom 3.5 to 10.2 mm in Empodisma minus. The leaves of Empodisma robustum are also longer, ranging from 2.2 to 7.55 mm, while the leaves range in length from 2.4 to 5.0 mm in Empodisma gracillimum and from 1.5 to 4.2 mm in Empodisma minus. The floral structures of Empodisma robustum are substantially longer than those of Empodisma minus. In contrast, the inflorescences of Empodisma gracillimum are smaller and more delicate then either Empodisma robustum or Empodisma minus (Fig. 6). The male spikelet of Empodisma robustum ranges from 6.8 to 9.0 mm in length, whereas Empodisma minus ranges from 3.9 to 8.0 mm and Empodisma gracillimum from 4.0 to 5.8 mm. The anthers in Empodisma robustum range from 1.9 to 2.5 mm in length, 1.2 to 2.0 in Empodisma minus, and 0.6 to 1.0 mm in Empodisma gracillimum. The female spikelets in Empodisma robustum ranges from 5.8 to 8.9 mm in length, 3.5 to 7.0 mm in Empodisma minus and 1.5 to 2.4 mm in Empodisma gracillimum. While few of the herbarium specimens that we examined had mature fruits, fruits from Empodisma robustum ranged from 2.6 to 2.8 mm in length, 2.3 to 3.0 mm in Empodisma minus, and 1.4 to 2.5 mm in Empodisma gracillimum.
Box plots illustrating patterns of morphological variation among the species of Empodisma. The box spans the interquartile range of the values in the variate. The middle 50% of the data lie within the box, with a line showing the median. The whiskers extend beyond the ends of the box as far as the minimum and maximum values.
The first PCoA axis accounted for 52.4 % of the variation in our sample and the second 23.4% (Fig. 7). The PCoA ordination separated Empodisma gracillimum primarily on the second axis but there was some overlap among the outliers of Empodisma robustum and Empodisma minus on the first axis. The greatest spread among the OTU’s was observed in Empodisma robustum and Empodisma gracillimum; this might reflect their taller more scrambling growth habit. With the exception of one outlier from Queensland, specimens of Empodisma minus are more tightly grouped. Several of the specimens of Empodisma minus were collected in the high mountains or in lowland bogs at more southerly latitudes. The stature of these plants may be more constrained by the harsh environments that they inhabit.
Principal coordinates ordination depicting patterns of overall similarity among the 74 OTEs that comprised our morphological sample. The first PC axis accounted for 58.3% of the variation in our sample and the second PC axis accounted for 23. 4% of the variation.
There is considerable uncertainty associated with estimating divergence times in Restionaceae. The fossil calibrations rely entirely upon microfossils and their affinities to extant genera are not clear. The divergence estimates presented here will undoubtedly be refined as more complete fossils are discovered. Nonetheless our preliminary findings suggest that Empodisma evolved in Australia during the mid Oligocene / early Miocene between 28–16 mya (Fig. 3). This was a time of warm, equitable environmental conditions.
The sequencingresults also differentiate the populations of Empodisma robustum from northern New Zealand from the Australian or New Zealand populations of Empodisma minus (Fig. 1), and this split was independently supported by each of the chloroplast DNA regions that we surveyed (Fig. 4). Empodisma robustum comprises a distinct evolutionary lineage united by six synapomorphies (Fig. 5). In contrast, the plants from mainland Australia and Tasmania are very similar to the diminutive lowland plants of Empodisma minus from southern New Zealand, each haplotype is distinguished only by a single mutation and the plants are found in similar habitats. The three genes that we sequenced were encoded in the chloroplast and are probably maternally inherited. Interestingly, there is no homoplasy in the data. Nonetheless, it is conceivable that the chloroplast gene tree is not compatible with the species tree. Gene convergence, introgression hybridization and/or incomplete lineage sorting could result in incompatible phylogenetic signals. A degree of reproductive isolation is necessary for these mutations to become fixed, which suggests Empodisma robustum and Empodisma minus have been reproductively isolated, perhaps since the Pleistocene. Within the northern haplotype, the sequences are identical. Although the sample is small, the absence of unique mutations (autapomorphies) suggests gene flow is unrestricted among the northern populations of Empodisma robustum. Historical rates of gene flow have traditionally been estimated indirectly from the number of fixed alleles in subpopulations relative to the total population (
Contrary to the observations of
The three species of Empodisma also have distinct ecological and distributional differences.
Ecology of Empodisma robustumEmpodisma robustum is restricted to the region north of 38° S latitude on the North Island of New Zealand. This phytogeographical boundary has long been recognized by New Zealand ecologists and marks the southernmost range of many species, most notably Agathis australis, but also bog associates such as Sporadanthus ferrugineus de Lange, Heenan & B.D.Clarkson, Dianella haematica Heenan & de Lange , Dracophyllum lessonianum A.Rich., Anzybas carsei (Cheeseman) D.L.Jones & M.A.Clem., and Lycopodiella serpentina (Kunze) B.Øllg. This region is the warmest in New Zealand and is rich in endemic species (
Empodisma robustum is a mid- to late-successional species of restiad raised bogs in the lowland zone of northern North Island (
Empodisma robustum is the key species in the fen–bog transition (
The development of raised bogs is constrained by a delicate water balance. They typically form in regions with moderate to high rainfall, cool summers, poor drainage, and isolation from flowing water (
Because the raised-bog surface is isolated from the influence of groundwater and surface runoff, plants receive their water and nutrients from rainfall. They typically have very low levels of plant nutrients, particularly of nitrogen and phosphorus (
Despite their saturated substrates, naturally occurring fires have been well documented in New Zealand peatlands (
Empodisma minus in New Zealand is also a mid- to late-successional wetland species. It dominates fens, blanket bogs, raised bogs, and pakihi heaths in coastal to alpine areas between 38° S latitude in the North Island and 48° S on Stewart Island, being particularly common in Westland and Southland. It is absent from Chatham Island. The vegetation is typically a dense, springy carpet of Empodisma minus, averaging 40 cm tall, associated with heath shrubs (Leptospermum scoparium, Dracophyllum oliveri Du Rietz, Dracophyllum prostratum Kirk), sedges (Baumea teretifolia, Baumea tenax (Hook.f.) S.T.Blake), the ferns Gleichenia dicarpa and Gleichenia microphylla R.Br., the tussock grass Chionochloa rubra Zotov, sundews (Drosera spp.), and Sphagnum cristatum moss.
Many of the ecological properties of Empodisma robustum also apply to Empodisma minus. For example, Empodisma minus forms peat via its cluster roots, although these are smaller and less dense than in Empodisma robustum. It is also a key species in the fen–bog transition, particularly Chionochloa rubra-dominated fens (
Empodisma minus resprouts after fire (
In Australia, Empodisma minus occurs in all states apart from Western Australia and Northern Territory, being concentrated in south-eastern Australia. It grows in similar habitats to New Zealand, i.e. fens and bogs, and seasonally or permanently inundated heaths, swamps and stream margins (
Empodisma gracillimum is endemic to Australia. It is restricted to the coastal plain from Perth southwards, and along the south coast from Augusta to Albany (
The three species of Empodisma form a well-supported clade. The clade diverged during the early Miocene, which was a period of equitable environmental conditions in Australia. A rapid succession of marine incursions, the onset of aridity in Australia, and origin of the Nullarbor Plain during the mid to late Miocene created barriers that isolated the southwest Australian endemic Empodisma gracillimum from the southeastern Australian Empodisma minus. Dispersal, colonization and speciation in New Zealand occurred more recently, coinciding with the uplift of the Southern Alps during the Pliocene and episodes of glaciation during the Pleistocene. Genetic, morphological and ecological evidence supports the separation of Empodisma minus into two species, Empodisma minus and Empodisma robustum. The split between Empodisma minus and Empodisma robustum is unambiguous and independently supported by the three cpDNA regions that we surveyed. Empodisma robustum is distinguished by six unique nucleotide substitutions and a 23-base duplication. It is a taller, more robust plant that is typically killed by fire and confined to lowland regions north of 38° S, whereas Empodisma minus is smaller, resprouts after fire, and occurs in alpine and lowland areas south of 38° S. The western Australian species Empodisma gracillimum emerges as sister to Empodisma minus and Empodisma robustum. It is geographically isolated and can be readily distinguished by its fine light green culms, shorter leaf sheaths and pedicellate female flowers. This last character appears to be a distinctive feature of the species.
TaxonomyEmpodisma minus (Hook.f.) L.A.S.Johnson & D.F.Cutler
Perennial herbs forming dense tangled masses, dioecious. Rhizomes stout up to 8.0 mm diam., covered with light brown, imbricate, scale-like sheaths and very thick tufts of brown hairs. Roots crowded, densely covered with persistent root hairs. Culms evergreen, hollow, dark to light green, profusely branching. Lamina reduced, awl-shaped, persistent, light green when young maturing dark brown, strongly reflexed from the leaf sheath. Leaf sheaths open, but overlapping and closely appressed, borne at short intervals, straw-coloured early in the season maturing dark brown, mouth ciliate with a tuft of woolly white hairs. Spikelets unisexual, borne in ultimate branch systems produced in second or third year, sessile or on short pedicels. Glumes imbricate; bracteoles lacking. Perianth segments 6, narrowly ovate almost hyaline. Male spikelets with 1–6 flowers, sessile to shortly pedicellate. Stamens 3, exserted beyond the perianth segments; filaments uniform; anthers linear oblong, dorsifixed, 1-celled, dehiscence along longitudinal slits, straw-coloured. Female spikelets solitary, each spikelet with 1– rarely 2 – flowers sessile to pedicillate. Ovary 1-celled; style branches 2 or 3, filiform, deciduous. Ovule solitary, pendulous. Fruit 1-seeded nut, ovoid with a thick and swollen base. 2n = 24. Fruit development is protracted with the seeds maturing in the following winter or early spring.
1 | Robust plants forming dense tangled thickets; culms > 1 mm in diameter at base of the plant; sheaths mostly >7.5 mm long; spikelets > 6.0 mm long | Empodisma robustum |
– | Slender and/or diminutive plants; culms mostly < 1 mm in diameter, sheaths mostly < 7.5 mm long; spikelets generally < 6.0 mm | 2. |
2 | Culms dark green, 18–80 cm in height (some plants rarely to 1.2 m in Eastern Australia); female spikelets sessile to shortly pedicellate | 2. Empodisma minus |
– | Culms light green, 55–130 cm in height; spikelets borne on pedicels up to 20 mm long | 3. Empodisma gracillimum |
(Fig. 8) New Zealand, Waikato, Hoe-O-Tainui, R. Mason, N.T. Moar 6750, 6/12/1958, CHR11159.
robustum describes the robust stature of Empodisma robustum.
Culms dark green, 38–139 cm in height (reportedly > 200 cm when supported by associated shrubs), 0.9–2.2 mm in diameter at the base, branching profusely. Leaf sheaths open, closely appressed, 5.2–21.0 mm in length, borne at intervals of 20.0–70.0 mm, light green to light brown early in the season maturing dark brown; mouth ciliate with a tuft of woolly white hairs. Lamina strongly reflexed from leaf sheath, 2.2–7.5 mm long, light green when young maturing dark brown. Spikelets brown; male spikelet 6.8–9.0 mm long, anthers 1.9–2.5 mm; female spikelet 5.8–8.9 mm; nutlets dark brown approximately 2.7 mm long. Flowering Aug.–Oct. See Figure 9.
Type of Empodisma robustum, N56 R. Mason, N.T. Moar 6750, 6/12/1958, CHR11159.
Morphological characteristics of Empodisma robustum. A Vegetative shoot with attached leaves and flowers (actual size) B Rhizomes with emerging vegetative shoots (2.5× actual size) C Vegetative shoot with attached pistillate flower D Pistillate flower with attached bracts E Gynoecium F Mature nut G Vegetative shoot with attached staminate spikelet H Staminate flower with attached bracts I Staminate flower. Scale bar = 1 mm.
Many herbarium specimens of Empodisma robustum include only the upper portion of the plant. These specimens may be difficult to distinguish from the larger specimens of Empodisma minus. Quality specimens should include a rhizome and the base of the culms, from which the distinguishing measurements are taken. Most collections of Empodisma robustum are either sterile or male, and the few females generally lack mature fruits. A chromosome count of 2n=24 was reported from plants collected at Moanatuatua Bog (
New Zealand, Lake Tangonge, H. Carse, H.B. Mathews, 25 Oct 1920, CHR295186; New Zealand, Moanatuatua Bog, W.F. Harris, 20 Nov 1951, CHR85625; New Zealand, Motutangi Swamp, T. Seymour, 15 July 1976, CHR287072; New Zealand, Tauhei, H. Carse, Aug 1925, CHR295191; New Zealand, Opuatia Bog, immature ♀ flowers J.T. Taylor, 27 July 1987, WAI8520; New Zealand, Torehape, not flowering, R.H. Chitty, WAI3280; New Zealand, Moanatuatua Bog, not flowering, R. Thompson, 3/77, WAI2099; New Zealand, Moanatuatua Bog, immature flower buds, H. Beaton 3/77, WAI2098; New Zealand, Kopouatai Peat Dome, immature flower buds, P.J. de Lange, 14 Mar 1988, WAI9008; New Zealand, ♂ in flower, R. Irving, M. Skinner, 12 Oct.1983, WAI 422; New Zealand, Tairua Ecological District, not flowering, B.R. Clarkson, 3 Feb 1998, WAI 16755; New Zealand, MoanatuatuaBog, not flowering, H.J. Beaton, 16 Aug1976, WAI 1100; New Zealand, Moanatuatua Bog, not flowering, K. Thompson 3/77, WAI 2100; New Zealand, Kaitaia, not flowering, W.F.B. Oliver, 26 Feb 1929, WELT19806; New Zealand, Ohaupo Swamp, ♂ flowers, T.F. Cheeseman, WELT19805; New Zealand, Ohaupo Swamp, not flowering, W. Petrie, WELT19804; New Zealand, Ohaupo Swamp, ♂ in flower, W. Petrie, WELT19803; New Zealand, Ohaupo Swamp, ♂ flowers, W. Petrie, WELT19802; New Zealand, Maitahi shrubland, not flowering, A.R Jamieson, AK231291; New Zealand, Rukuhia Swamp ♂ and ♀ plants in flower, L.M. Cranwell 18/34, AK109372; New Zealand, Lake Ohia, ♂ flowers, R. Cooper, R. Mason, N. Moar 1 Aug 1949, AK35820; New Zealand, Torehape Peat Dome ♂ flowers, A.E. Wright 10576, AK215859; New Zealand, Whangamarino Swamp, ♂ flowers, plants up to 1.2 m tall, E.K. Cameron 8839; AK234026; New Zealand, Kaihu Valley, ♂ flowers, A.R. Jamieson 30 Oct 1999, AK286616; New Zealand, Tomarata Lakes, not flowering, M.E. Young 20 March 2007, AK299780; New Zealand, Moanatuatua Peat Reserve, Rukuhia, ♂ flowers, F.J. Newhook July 1979, AK304253; New Zealand, Rukuhia Swamp, separate plants with ♂ and ♀ flowers, L.M. Cranwell 18/34, AK109373; New Zealand, Mangawhai Black Swamp, immature ♀flowers, M.E. Young 18 July 1999, AK239846; New Zealand, Mercer Swamp, P. Hynes 15 Feb 1964, AK101004; New Zealand, Tomarata Lakes, with few ♀ flowers, M.E. Young 20 March 2007, AK299780; New Zealand, Lake Ohia, ♂ flowers, A.E. Wright 10554, AK232056; New Zealand, Lake Ohia, ♀ flowers and fruits, A.E. Wright 10555, AK232055; New Zealand, Tokerau Beach, M.E. Young, L.J. Forester 17 Oct 2006, AK306920; New Zealand, Lake Ohia, ♂ flowers, J.E. Braggins 87/87A, AK304249.
The acronym for the University of Waikato herbarium was recently changed from WAI to WAIK. We cited the older WAI acronym which appeared on specimen labels that we studied.
New Zealand endemic ranging from North Cape southwards to approximately 38° S latitude.
Empodisma robustum is restricted to ombrotrophic raised peat bogs where it often coexists with Sporadanthus ferrugineus, fens and gumland heathland peats. Locally abundant, but populations becoming fragmented by intensive land use.
Widespread drainage and conversion to pasture has dramatically reduced the extent of raised peat bogs in Northland and Waikato. This unique ecosystem is severely fragmented and provides habitat for a number of rare plant species such as Sporadanthus ferrugineus (
http://species-id.net/wiki/Empodisma_minus
New Zealand, near Nelson, Bidwell no. 84, K000441989; (Fig. 10; designated by
High resolution photograph of the lectotype of Empodisma minus (Hook.f.) L.A.S. Johnson & D.F.Cutler. Reproduced with the consent of the Royal Botanic Gardens, Kew, © The Board of Trustees of the Royal Botanic Gardens.
minus describes the small stature of Empodisma minus.
Culms dark green, 12–81 cm in height, 0.7–1.3 mm in diameter, branching profusely. Leaf sheaths closely appressed, 3.5–10.2 mm in length, borne at short intervals 15.0–48.0 mm; light green to light brown early in the season maturing dark brown; mouth ciliate with a prominent tuft of woolly white hairs in New Zealand specimens, spare or lacking in Australian specimens. Lamina strongly reflexed from leaf sheath, 1.5–4.2 mm long, persistent light green when young maturing dark brown. Spikelets brown, male spikelet 3.9–8 mm long, anthers 1.2–2.0 mm long; female spikelet 3.5–7.0 mm long; nutlets dark brown approximately 2.6 mm long. 2n = 24. Flowering Aug.– Apr. [Fig. 11; see also illustration in
Morphological characteristics of Empodisma minus. A Vegetative shoot with attached leaves and flowers (2.5× actual size) B Rhizomes with emerging vegetative shoots (2.5× actual size) C Vegetative shoot with attached pistillate flower D Pistillate flower with attached bracts E Gynoecium F Mature nut G Vegetative shoot with attached staminate spikelet H Staminate flower with attached bracts I Staminate flower. Scale bar = 1 mm.
Morphologically similar to small forms of Empodisma robustum distinguished by its smaller stature (though plants from 0.4 to 1 m tall are noted from eastern Australia), more delicate culms and smaller spikelets. Most collections are sterile or male, and the few females generally lack mature fruits. Chromosome counts of 2n=24 were reported from plants collected from the NW slope of Mt. Ruapehu and three counts from NSW (
Australia, Tasmania, near Margate, ♂ flowers, D.A. & A. Ratkowsky 1474, CHR303032, Australia, Queensland, Moreton Island, ♂ flowers, L. Durrington 1114 & S. Levine, CHR272564, Australia, Tasmania, Newdegate Pass, not in flower, T. Dobson 77107, CHR313744; New Zealand, S. Westland, ♂ flowers, G.C. Kelly, Oct 1966, CHR177206; New Zealand, Rahu Saddle, with few ♀ flowers, E.J. Godley, 1 July 1958, CHR108315; New Zealand, Bell Hill Plains, ♂ flowers, J. Clarke 1 Feb 1969, CHR189013; New Zealand, Ngamatea, ♂ flowers, N.J. Moar, 12 Jan 1949, CHR70144; New Zealand, Waikareiti, ♂ flowers, A.P. Druce, Feb 1968, CHR180674; New Zealand, Silica Springs Track, ♂ flowers G. Rennison, A61/36, , CHR535708; New Zealand, Makerikeri tarns, ♂ flowers, A.P. Druce, Nov 1973, CHR260376; New Zealand, Tussock Creek, ♂ flowers, L.B. Moore, 28 July 1968, CHR188099; New Zealand, Mokoreta, with few ♀ flowers, W.R. Sykes 41/94, CHR497058; New Zealand, Bayswater Bog, not flowering, B.R. Clarkson19 Feb .2009, CHR605146; New Zealand, Awarua Bay, not flowering, P.N. Johnson 653, CHR437892; New Zealand, West Cape, A.F. Mark, 5 Feb 1972, CHR218694; New Zealand, Coal Creek, ♂ flowers; I. Payton, 13 Sept 1976, CHR520808; New Zealand, Mt. Rockport, not flowering, I.A. McNew, 31 July 1942, CHR35234; New Zealand, Lake Sylvester, ♂ flowers, R. Melville 5915, CHR142781; New Zealand, Lake Sylvester, ♀ flowers, R. Mason & N. Moar 4658, CHR95709; New Zealand, Bealey spur, with immature ♀ flowers, P. Douglas 26 Nov 1979, CHR362302; New Zealand, Lake Tennyson, ♂ flowers, M.J.A. Simpson 6315, CHR22759; New Zealand, Patterson Inlet, ♂ flowers, L.J. Dumbleton & E. Edgar, CHR182509; New Zealand, ♂ flowers, Fosberg, Feb 1949; CHR30378; New Zealand, Kaitangata, plants with ♀ and ♂ flowers, R. Mason & N.T. Moar 953, CHR 75833; New Zealand, Kapuka, ♀ flowers, W.H. Harbond 20 Nov 1968, CHR183615.
Widely distributed in Tasmania and all mainland Australian States except Western Australia and the Northern Territory; in New Zealand extending north to approximately 38°S latitude.
Locally abundant in seasonally or permanently inundated wetlands, heathlands, fens and peat bogs from sea level to alpine
Not threatened.
http://species-id.net/wiki/Empodisma_gracillimum
(Fig. 12; fide BG Briggs xi.1998), Nouvelle-Hollande, Riv. des cygnes, Preiss JA 1711, 1843. P00748711; Nouvelle-Hollande, Riv. des cygnes, Preiss JA 1714, 1843. P00748712.
High resolution photograph of a syntype of Empodisma gracillimum (F.Muell.) L.A.S. Johnson & D.F.Cutler.
gracillimum describes the slender culms of Empodisma gracillimum.
(illustrated in
Though it approaches Empodisma robustum in height, Empodisma gracillimum is readily distinguished by its more delicate light green culms and shorter leaf sheaths. The male and female spikelets of Empodisma gracillimum are smaller than either Empodisma robustum or Empodisma minus. The female spikelets are solitary and distinctly pedicillate; this character may be a synapomorphy for the species.
Western Australia, Denmark, ♂ and ♀ flowers, B.G. Briggs 8449 & L.A.S. Johnson, CHR525963; Western Australia, 4.4 km east Watershed Road, ♂ flowers, A.R. Annels, R.W. Hearn 5112, PERTH04219031; Australia, 4.4 km east Watershed Road, fruits, A.R. Annels, R.. Hearn 5111, PERTH04128567; Western Australia, Denmark, ♀ flowers, B.G. Briggs 8449 & L.A.S. Johnson, PERTH01586645; Western Australia, S. of junction with Brockman Highway, ♀ flowers D. Bright, C. Godden & T. Annels SC72.9 PERTH04723732; Western Australia, London Forest Block, 2km S of Mountain Road along Renzo Road, ♂ flowers, R.J. Cranfield & B.G. Ward WFM53, PERTH07102399; Western Australia, Torndirrup National Park, ♂ flowers, G.J. Keighery 8805, PERTH02182831; Western Australia, Darling, ♂ flowers, B.G. Briggs 9330, PERTH06173853; Western Australia, 800 m E along O’Byrene Road from intersection of Commonage Road, ♂ flowers, N. Casson & T. Annels SC32.9, PERTH04741110; Western Australia, Walpole-Nornalup National Park, ♀ flowers, A.R. Annels ARA1580, PERTH05466172; Western Australia, Walpole, R.J. Cranfield 10897, PERTH04638530; Western Australia, 600 m S of Brockman Highway on Beck Road, ♂ flowers, N. Casson, P. Ellery & C. McChesney SC74.8, PERTH04723775; Western Australia, 400 m E of Blackwood and Great North Road, ♂ flowers, R. Davis 7680, PERTH05139317; Western Australia, WA, N. Casson & D. Bright SC106.2, PERTH04749677; Western Australia, S.E. Witchcliff, ♂ flowers, G.J. Keighery 16277, PERTH06330266; Western Australia, NE of Albany, E.M. Sandiford & D.A. Rathbone 1372, PERTH07926855.
Endemic to western Australia on the coastal plain south of Perth extending along the south coast from Augusta to Albany.
Grows on peat or sandy nutrient-poor soils. Locally abundant in seasonally or permanently inundated wetlands, swamps and stream margins.
We thank the curators of the Western Australia Herbarium (PERTH), Auckland Museum (AK), Te Papa-The National Museum of New Zealand (WELT), Waikato University Herbarium (WAIK) and Allan Herbarium (CHR) for the assistance with loans. The type of Empodisma minus was reproduced with the consent of the Royal Botanic Gardens, Kew, © The Board of Trustees of the Royal Botanic Gardens and Empodisma gracillimum the Muséum National d’Histoire Naturelle (MNHN). Rebecca Wagstaff provided the beautifully rendered illustrations of Empodisma minus and Empodisma robustum. Scott Bartlam and Jane Cruickshank helped with field and technical assistance. Earlier versions of this manuscript benefited greatly from the constructive suggestions provided by Ilse Breitwieser, David Glenny, Peter Heenan, Peter Linder and an anonymous reviewer. We would also like to acknowledge the careful editing of Christine Bezar and Anne Austin. This research was supported by the Ministry of Science and Innovation through the Defining New Zealand’s Land Biota and the Restoring Wetlands contracts.
Voucher specimens listing country of origin, literature citations, Allan Herbarium accession numbers for new sequences, their collection locality, DNA accession number, and GenBank accession numbers.
Specie | Country | Literature citation | Locality | Herbarium accession | DNA accession | matK | rbcL | trnL |
---|---|---|---|---|---|---|---|---|
Alexgeorgea subterranean Carlquist | Australia |
|
NSW 437369 | GQ409034 | GQ408918 | GQ408988 | ||
Anarthria polyphylla Nees | Australia |
|
NSW 391527 | DQ257498 | AF148760 | AF148720 | ||
Apodasmia brownii (Hook.f.) B.G.Briggs & L.A.S.Johnson | Australia |
|
NSW 494422 | — | GQ408919 | — | ||
Apodasmia chilensis (Gay) B.G.Briggs & L.A.S.Johnson | Chile |
|
Los Lagos, Mehuín | CHR513924 | JX154568 | AF307923 | JX154570 | |
Apodasmia similis (Edgar) B.G.Briggs & L.A.S.Johnson | New Zealand |
|
Westland, Wanganui River | CHR517317 | JX154569 | AF307924 | JX154571 | |
Baloskion tetraphyllum (Labill.) B.G.Briggs & L.A.S.Johnson | Australia |
|
NSW365050 | DQ257501 | AF148761 | AF148721 | ||
Calorophus elongatus Labill. | Australia |
|
NSW 264835 | GQ409036 | DQ257502 | AF148725 | ||
Calorophus erostris (C.B.Clarke) L.A.S.Johnson & B.G.Briggs | Australia |
|
NSW 264698 | GQ409036 | GQ408930 | GQ408999 | ||
Chaetanthus aristatus (R.Br.) B.G.Briggs & L.A.S.Johnson | Australia |
|
NSW 261929 | DQ257508 | AF148782 | AF148743 | ||
Chordifex crispatus (R.Br.) B.G.Briggs & L.A.S.Johnson | Australia |
|
NSW 401500 | DQ257510 | GQ408923 | GQ408922 | ||
Chordifex dimorphus (R.Br.) B.G.Briggs | Australia |
|
NSW 270162 | — | AF148763 | AF148723 | ||
Chordifex fastigiatus (R.Br.) B.G.Briggs | Australia |
|
NSW 270160 | — | AF148791 | AF148752 | ||
Chordifex hookeri (D.I.Morris) B.G.Briggs, | Australia |
|
NSW 264839 | — | AF148762 | AF148722 | ||
Coleocarya gracilis S.T.Blake | Australia |
|
NSW 401500 | GQ409023 | AF148769 | AF148730 | ||
Dapsilanthus ramosus (R.Br.) B.G.Briggs & L.A.S.Johnson | Australia |
|
NSW 338881 | — | AF148780 | AF148741 | ||
Desmocladus castaneus B.G.Briggs & L.A.S.Johnson | Australia |
|
NSW 423447 | DQ257511 | AF148770 | AF148731 | ||
Dielsia stenostachya (W.Fitzg.) B.G.Briggs & L.A.S.Johnson | Australia |
|
NSW 391321 | — | AF148771 | AF148732 | ||
Empodisma gracillimum (F.Muell.) L.A.S. Johnson & D.F.Cutler | Australia | This paper | Western Australia, Denmark | CHR 525963 | 9.14 | JX129074 | JX129095 | JX129133 |
Empodisma gracillimum (F.Muell.) L.A.S. Johnson & D.F.Cutler | Australia | This paper | Western Australia, Caldyannup land system | PERTH 07102399 | 12.1 | JX129076 | JX129095 | JX129132 |
Empodisma gracillimum (F.Muell.) L.A.S. Johnson & D.F.Cutler | Australia | This paper | Western Australia, NE of Albany | PERTH 046385530 | 12.2 | JX129075 | JX129097 | JX129131 |
Empodisma minus (Hook.f.) L.A.S.Johnson & D.F.Cutler | Australia | This paper | Tasmania, Central Plateau, second lagoon | CHR 585759 | 7.45 | JX129080 | JX129101 | JX129116 |
Empodisma minus (Hook.f.) L.A.S.Johnson & D.F.Cutler | Australia | This paper | Victoria, Mt. Buffalo | CHR 607930 | 9.15 | JX129090 | JX129111 | JX129117 |
Empodisma minus (Hook.f.) L.A.S.Johnson & D.F.Cutler | New Zealand | This paper | Waipapa EA, Pureora | CHR 605145 | 9.04 | JX129087 | JX129108 | JX129119 |
Empodisma minus (Hook.f.) L.A.S.Johnson & D.F.Cutler | New Zealand | This paper | Lookout Range | CHR 605066 | 8.09 | JX129085 | JX129106 | JX129120 |
Empodisma minus (Hook.f.) L.A.S.Johnson & D.F.Cutler | New Zealand | This paper | Herangi Range, top | CHR 596548 | 8.08 | JX129084 | JX129105 | JX129121 |
Empodisma minus (Hook.f.) L.A.S.Johnson & D.F.Cutler | New Zealand | This paper | Taramoa | CHR 605065 | 8.01 | JX129081 | JX129102 | JX129122 |
Empodisma minus (Hook.f.) L.A.S.Johnson & D.F.Cutler | New Zealand | This paper | Stewart Island, Mason Bay | CHR 605074 | 7.40 | JX129077 | JX129098 | JX129123 |
Empodisma minus (Hook.f.) L.A.S.Johnson & D.F.Cutler | New Zealand | This paper | Pukerau Red Tussock Reserve | CHR 605847 | 9.05 | JX129088 | JX129109 | JX129118 |
Empodisma robustum Wagstaff & Clarkson sp. nov. | New Zealand | This paper | Whangamarino | CHR 605067 | 11.01 | JX129091 | JX129112 | JX129124 |
Empodisma robustum Wagstaff & Clarkson sp. nov. | New Zealand | This paper | Opuatia | CHR 605068 | 7.44 | JX129079 | JX129100 | JX129129 |
Empodisma robustum Wagstaff & Clarkson sp. nov. | New Zealand | This paper | Moanatuatua Swamp | CHR 605165 | 9.06 | JX129089 | JX129110 | JX129125 |
Empodisma robustum Wagstaff & Clarkson sp. nov. | New Zealand | This paper | Northland, Tangonge | CHR 605064 | 8.03 | JX129082 | JX129103 | JX129128 |
Empodisma robustum Wagstaff & Clarkson sp. nov. | New Zealand | This paper | Lake Tomarata | CHR 605069 | 7.43 | JX129078 | JX129099 | JX129130 |
Empodisma robustum Wagstaff & Clarkson sp. nov. | New Zealand | This paper | Kaimaumau | CHR 605063 | 8.06 | JX129083 | JX129104 | JX129127 |
Empodisma robustum Wagstaff & Clarkson sp. nov. | New Zealand | This paper | Torehape Bog | CHR 605167 | 9.01 | JX129086 | JX129107 | JX129126 |
Eurychorda complanata (R.Br.) B.G.Briggs & L.A.S.Johnson | Australia |
|
NSW 264949 | DQ257514 | AF148790 | AF148751 | ||
Harperia lateriflora W.Fitzg. | Australia |
|
NSW 423455 | GQ409020 | AF148776 | AF148737 | ||
Hopkinsia adscendens B.G.Briggs & L.A.S.Johnson | Australia |
|
NSW 364372 | DQ257519 | AF148727 | AF148738 | ||
Hypolaena exsulca R.Br. | Australia |
|
NSW 364832 | — | GQQ408927 | — | ||
Hypolaena grandiuscula F.Muell. | Australia |
|
NSW 714757 | — | — | GQ408962 | ||
Hypolaena pubescens (R.Br.) Nees | Australia |
|
NSW 714454 | GQ409046 | — | GQ409963 | ||
Hypolaena robusta Meney & Pate | Australia |
|
NSW 714451 | — | — | GQ408964 | ||
Kulinia eludens B.G.Briggs & L.A.S.Johnson | Australia |
|
NSW 391535 | — | AF148778 | AF148739 | ||
Lepidobolus chaetocephalus Benth. | Australia |
|
NSW 364813 | — | AF148779 | AF148740 | ||
Leptocarpus tenax (Labill.) R.Br. | Australia |
|
NSW 264954 | GQ409039 | AF148781 | AF148742 | ||
Lepyrodia glauca (Nees) F.Muell. | Australia |
|
NSW 423726 | — | AF148785 | AF148746 | ||
Loxocarya gigas B.G.Briggs & L.A.S.Johnson | Australia |
|
NSW 364738 | — | AF148786 | AF148747 | ||
Lyginia barbata R.Br. | Australia |
|
NSW 391339 | DQ257523 | AF148787 | AF148748 | ||
Meeboldina coangustata (Nees) B.G.Briggs & L.A.S.Johnson | Australia |
|
NSW 261610 | GQ409043 | AF148284 | AF148745 | ||
Melanostachya ustulata (F.Muell. ex Ewart & Sharman) B.G.Briggs & L.A.S.Johnson | Australia |
|
NSW 232599 | GQ409035 | AF148788 | AF148749 | ||
Restio distichus Rottb. | Africa |
|
Linder, Hardy and Moline 7327 | AY881540 | AY881467 | AY881613 | ||
Sporadanthus ferrugineus de Lange, Heenan & B.D.Clarkson | New Zealand | This paper | South Auckland; Moanatuatua Swamp | CHR 605163 | 9.10 | JX129093 | JX129114 | JX129135 |
Sporadanthus ferrugineus de Lange, Heenan & B.D.Clarkson | New Zealand | This paper | South Auckland; Kopuati | CHR 604580 | 8.56 | JX129092 | JX129113 | JX129134 |
Sporadanthus gracilis (R.Br.) B.G.Briggs & L.A.S.Johnson | Australia |
|
NSW 270154 | GQ409027 | DQ257526 | GQ409013 | ||
Sporadanthus tasmanicus (Hook.f.) B.G.Briggs & L.A.S.Johnson | Australia |
|
NSW 264956 | GQ409028 | AF148793 | AF148754 | ||
Sporadanthus traversii F.Muell. | New Zealand | This paper | Chatham Islands; Rakautahi | CHR 605164 | 9.08 | JX129094 | JX129115 | JX129136 |
Taraxis grossa B.G.Briggs & L.A.S.Johnson | Australia |
|
NSW 270154 | GQ409027 | DQ257526 | GQ409013 | ||
Tremulina tremula B.G.Briggs & L.A.S.Johnson | Australia |
|
NSW 264956 | GQ409028 | AF148793 | AF148754 | ||
Tyrbastes glaucescens B.G.Briggs & L.A.S.Johnson | Australia |
|
NSW 261641 | GQ409037 | AF148795 | AF148756 | ||
Winifredia sola L.A.S.Johnson & B.G.Briggs | Australia |
|
NSW 713239 | GQ409021 | AF148796 | AF148758 |