Register      Login
Australian Systematic Botany Australian Systematic Botany Society
Taxonomy, biogeography and evolution of plants
RESEARCH ARTICLE (Open Access)

Morphological and molecular evidence for major re-circumscriptions in and eight new species of Melichrus R.Br. (Ericaceae subfam. Epacridoideae) in eastern Australia

Helen T. Kennedy https://orcid.org/0000-0003-4856-5169 A B * , Ian R. H. Telford https://orcid.org/0000-0002-3570-0053 A , Darren Crayn https://orcid.org/0000-0001-6614-4216 C D , Jeremy J. Bruhl https://orcid.org/0000-0001-9112-4436 A and Rose L. Andrew https://orcid.org/0000-0003-0099-8336 A
+ Author Affiliations
- Author Affiliations

A Botany and N.C.W. Beadle Herbarium, School of Environmental and Rural Science, University of New England, Armidale, NSW 2351, Australia.

B Australian National Herbarium, Centre for Australian National Biodiversity Research (a joint venture between Parks Australia and CSIRO), GPO Box 1700, Canberra, ACT 2601, Australia.

C Australian Tropical Herbarium, James Cook University, Cairns, Qld 4870, Australia.

D Centre for Tropical Environmental Sustainability Science, James Cook University, Cairns, Qld 4870, Australia.

* Correspondence to: helen.kennedy@csiro.au

Handling Editor: Russell Barrett

Australian Systematic Botany 38, SB24031 https://doi.org/10.1071/SB24031
Submitted: 2 September 2024  Accepted: 2 March 2025  Published: 16 April 2025

© 2025 The Author(s) (or their employer(s)). Published by CSIRO Publishing. This is an open access article distributed under the Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License (CC BY-NC-ND)

Abstract

The genus Melichrus R.Br. has received very little taxonomic attention and treatments have largely disagreed on species delimitation. The eastern Australian clade of Melichrus was last revised in 1958. Over 60 years later we present new, in-depth evidence for species delimitation. A morphological dataset (90 individuals from 68 populations, scored for 26 characters) was analysed using NMDS ordination with Bayesian Inference cluster modelling (mclust) and UPGMA hierarchical clustering to detect morphological discontinuities in the genus that may indicate species boundaries. Discontinuities in the morphological analyses were compared with those apparent in clustering (principal component analysis, SplitsTree Neighbour-Net, STRUCTURE and conStruct) and statistical analyses (HE, HO, FIS and FST) of a DArTseq SNP dataset of 548 samples from 110 populations of Melichrus. These new lines of evidence form the basis of detailed recommendations for a revised species taxonomy of Melichrus including the description of eight new species, recircumscription of previously described species and the correction of a longstanding nomenclatural misapplication.

Keywords: DArTseq, Epacridoideae, Ericaceae, Melichrus, morphometrics, phenetics, reduced representation, species delimitation, taxonomy.

References

Barker WR (2005) Standardising informal names in Australian publications. Australian Systematic Botany Society Newsletter 122, 11-12.
| Google Scholar |

Bartling FG (1830) ‘Natural ordines plantarum eorumque characteres et affinitates adjecta generum enumeration.’ (Sumtibus Dieterichianis: Gottingen, German Confederation) [In Latin]

Bentham G (1868) ‘Flora Australiensis: A description of the plants of the Australian Territory. Vol. IV. Stylidieae to Pedalineae.’ (Reeve: London, UK)

Bentham G, Hooker JD (1876) ‘Genera plantarum: ad exemplaria imprimis in Herberiis Kewensibus servata definita.’ (A. Black: London, UK) [In Latin]

Bradburd GS, Coop GM, Ralph PL (2018) Inferring continuous and discrete population genetic structure across space. Genetics 210, 33-52.
| Crossref | Google Scholar | PubMed |

Brown R (1810) ‘Prodromus florae Novae Hollandiae et Insulae Van-Diemen, exhibens characteres plantarum.’ (Johnson & Co: London, UK) [In Latin]

Bryant D, Moulton V (2004) Neighbor-Net: an agglomerative method for the construction of phylogenetic networks. Molecular Biology and Evolution 21, 255-265.
| Crossref | Google Scholar | PubMed |

Centre for Australian National Biodiversity Research (2018) Botanical Districts of Australia. (Australian National Herbarium) Available at https://www.anbg.gov.au/cpbr/anhsir/anhsir-manual/botanical-districts.html [Verified 31 March 2025]

Carstens BC, Pelletier TA, Reid NM, Satler JD (2013) How to fail at species delimitation. Molecular Ecology 22, 4369-4383.
| Crossref | Google Scholar | PubMed |

Carvajal DE, Loayza AP, Rios RS, Gianoli E, Squeo FA (2017) Population variation in drought-resistance strategies in a desert shrub along an aridity gradient: Interplay between phenotypic plasticity and ecotypic differentiation. Perspectives in Plant Ecology, Evolution and Systematics 29, 12-19.
| Crossref | Google Scholar |

Cavanilles AJ (1797) ‘Icones et descriptiones plantarum, quae aut sponte in Hispania crescunt, aut in hortis hospitantur.’ (Regia typographia: Madrid, Kindom of Spain) [In Latin]

Collins TL, Schmidt-Lebuhn AN, Andrew RL, Telford IRH, Bruhl JJ (2022) There’s gold in them thar hills! Morphology and molecules delimit species in Xerochrysum (Asteraceae; Gnaphalieae) and reveal many new taxa. Australian Systematic Botany 35, 120-185.
| Crossref | Google Scholar |

Crayn DM, Hislop M, Puente-Lelièvre C (2020) A phylogenetic recircumscription of Styphelia (Ericaceae, Epacridoideae, Styphelieae). Australian Systematic Botany 33, 137-168.
| Crossref | Google Scholar |

Dayrat B (2005) Towards integrative taxonomy. Biological Journal of the Linnean Society 85, 407-415.
| Crossref | Google Scholar |

De Candolle AP (1839) ‘Prodromus Systematis Naturalis Regni Vegetabilis.’ (Treuttel and Wurtz: Paris, France) [In Latin]

Denham SS, Brignone NF, Johnson LA, Pozner RE (2019) Using integrative taxonomy and multispecies coalescent models for phylogeny reconstruction and species delimitation within the “Nastanthus–Gamocarpha” clade (Calyceraceae). Molecular Phylogenetics and Evolution 130, 211-226.
| Crossref | Google Scholar | PubMed |

de Queiroz K (2007) Species concepts and species delimitation. Systematic Biology 56, 879-86.
| Crossref | Google Scholar | PubMed |

Druce GC (1917) Nomenclatural Notes: chiefly African and Australian. Botanical Society and Exchange Club of the British Isles Report for 1916 4, 635.
| Google Scholar |

Drude O (1887) Epacridaceae. In ‘Die Natiirlichen Pflanzenfamilie. Vol. Teil 4, Abt. 1–2’. (Eds A Engler, K Prantl) pp. 66–79. (W. Engelmann: Leipzig, German Empire) [In German]

Edwards DL, Knowles LL (2014) Species detection and individual assignment in species delimitation: can integrative data increase efficacy? Proceedings of the Royal Society of London – B. Biological Sciences 281(1777), 20132765.
| Crossref | Google Scholar | PubMed |

Ewart K, Lo N, Ogden R, Joseph L, Ho S, Frankham G, Eldridge M, Schodde R, Johnson R (2020) Phylogeography of the iconic Australian red-tailed black-cockatoo (Calyptorhynchus banksii) and implications for its conservation. Heredity 125, 167.
| Crossref | Google Scholar |

Field B (Ed.) (1825) ‘Geographical Memoirs on New South Wales; by Various Hands.’ (John Murray: London, UK) Available at http://gutenberg.net.au/ebooks13/1304421h.html

Fraley C, Raftery AE (2002) Model-based clustering, discriminant analysis, and density estimation. Journal of the American Statistical Association 97, 611-631.
| Crossref | Google Scholar |

Fujita MK, Leaché AD, Burbrink FT, McGuire JA, Moritz C (2012) Coalescent-based species delimitation in an integrative taxonomy. Trends in Ecology & Evolution 27, 480-488.
| Crossref | Google Scholar | PubMed |

Funk SM, Guedaoura S, Juras R, Raziq A, Landolsi F, Luís C, Martínez AM, Musa Mayaki A, Mujica F, Oom MdM, Ouragh L, Stranger Y-M, Vega-Pla JL, Cothran EG (2020) Major inconsistencies of inferred population genetic structure estimated in a large set of domestic horse breeds using microsatellites. Ecology and Evolution 10, 4261-4279.
| Crossref | Google Scholar | PubMed |

Gower JC (1971) A general coefficient of similarity and some of its properties. Biometrics 27, 857-871.
| Crossref | Google Scholar |

Gruber B, Unmack PJ, Berry OF, Georges A (2018) dartR: an R package to facilitate analysis of SNP data generated from reduced representation genome sequencing. Molecular Ecology Resources 18, 691-699.
| Crossref | Google Scholar | PubMed |

Hausdorf B, Hennig C (2010) Species delimitation using dominant and codominant multilocus markers. Systematic Biology 59(5), 491-503.
| Crossref | Google Scholar | PubMed |

Hörandl E, Stuessy TF (2010) Paraphyletic groups as natural units of biological classification. Taxon 59, 1641-1653.
| Crossref | Google Scholar |

Hosegood J, Humble E, Ogden R, de Bruyn M, Creer S, Stevens GMW, Abudaya M, Bassos-Hull K, Bonfil R, Fernando D, Foote AD, Hipperson H, Jabado RW, Kaden J, Moazzam M, Peel LR, Pollett S, Ponzo A, Poortvliet M, Salah J, Senn H, Stewart JD, Wintner S, Carvalho G (2020) Phylogenomics and species delimitation for effective conservation of manta and devil rays. Molecular Ecology 29, 4783-4796.
| Crossref | Google Scholar | PubMed |

Huson DH, Bryant D (2006) Application of phylogenetic networks in evolutionary studies. Molecular Biology and Evolution 23, 254-267.
| Crossref | Google Scholar | PubMed |

Janes JK, Miller JM, Dupuis JR, Malenfant RM, Gorrell JC, Cullingham CI, Andrew RL (2017) The K=2 conundrum. Molecular Ecology 26, 3594-3602.
| Crossref | Google Scholar | PubMed |

Kennedy HT, Telford IRH, Crayn DM, Bruhl JJ (2020) Validation of two informally named species of Melichrus (Ericaceae: Epacridoideae) from north-eastern New South Wales. Telopea 23, 187-196.
| Crossref | Google Scholar |

Kilian A, Wenzl P, Huttner E, Carling J, Xia L, Blois H, Caig V, Heller-Uszynska K, Jaccoud D, Hopper C, Aschenbrenner-Kilian M, Evers M, Peng K, Cayla C, Hok P, Uszynski G (2012) Diversity Arrays Technology: a generic genome profiling technology on open platforms. In ‘Data Production and Analysis in Population Genomics: Methods and Protocols’. (Eds F Pompanon, A Bonin) pp. 67–89. (Humana Press: Totowa, NJ, USA)

Kopelman NM, Mayzel J, Jakobsson M, Rosenberg NA, Mayrose I (2015) Clumpak: a program for identifying clustering modes and packaging population structure inferences across K. Molecular Ecology Resources 15, 1179-1191.
| Crossref | Google Scholar | PubMed |

Kron KA, Judd WS, Stevens PF, Crayn DM, Anderberg AA, Gadek PA, Quinn CJ, Luteyn JL (2002) Phylogenetic classification of Ericaceae: molecular and morphological evidence. Botanical Review 68, 335-423.
| Crossref | Google Scholar |

Kruskal JB (1964) Multidimensional scaling by optimizing goodness of fit to a nonmetric hypothesis. Psychometrika 29, 1-27.
| Crossref | Google Scholar |

Lawson DJ, van Dorp L, Falush D (2018) A tutorial on how not to over-interpret STRUCTURE and ADMIXTURE bar plots. Nature Communications 9, 3258.
| Crossref | Google Scholar | PubMed |

Linck E, Battey CJ (2019) Minor allele frequency thresholds strongly affect population structure inference with genomic data sets. Molecular Ecology Resources 19, 639-647.
| Crossref | Google Scholar | PubMed |

Mace GM (2004) The role of taxonomy in species conservation. Philosophical Transactions of the Royal Society of London – B. Biological Sciences 359, 711-719.
| Crossref | Google Scholar | PubMed |

McCusker A (1999) Glossary. In ‘Flora of Australia. Vol. 1.’ (Eds AE Orchard, HS Thompson) pp. 585–636. (Australian Government Publishing Service: Canberra, ACT, Australia)

Meudt HM, Thorsen MJ, Prebble JM (2020) Taxonomic revision of the Myosotis australis group (Boraginaceae) native to Australia, New Zealand and New Guinea. Australian Systematic Botany 33, 477-524.
| Crossref | Google Scholar |

Mijangos JL, Gruber B, Berry O, Pacioni C, Georges A (2022) dartR v2: an accessible genetic analysis platform for conservation, ecology and agriculture. Methods in Ecology and Evolution 13, 2150-2158.
| Crossref | Google Scholar |

Nauheimer L, Weigner N, Joyce E, Crayn D, Clarke C, Nargar K (2021) HybPhaser: a workflow for the detection and phasing of hybrids in target capture datasets. Applications in Plant Sciences 9, e11441.
| Crossref | Google Scholar | PubMed |

New South Wales Department of Planning, Industry and Environment (2021) ‘Mount Kaputar National Park Plan of Management.’ (Environment, Energy and Science, Department of Planning, Industry and Environment: Sydney, NSW, Australia)

Orchard AE (1999) A history of systematic botany in Australia. In ‘Flora of Australia, Vol. 1’. (Eds AE Orchard, HS Thompson) pp. 11–103. (Australian Biological Resources Study: Canberra, ACT, Australia; and CSIRO Publishing: Melbourne, Vic., Australia)

Padial JM, Miralles A, De la Riva I, Vences M (2010) The integrative future of taxonomy. Frontiers in Zoology 7, 16.
| Crossref | Google Scholar | PubMed |

Paterson BR (1958) Revision of the genus Melichrus R.Br. (Epacridaceae). Proceedings of the Linnean Society of New South Wales 82, 303-313.
| Google Scholar |

Pembleton LW, Cogan NO, Forster JW (2013) StAMPP: an R package for calculation of genetic differentiation and structure of mixed-ploidy level populations. Molecular Ecology Resources 13, 946-952.
| Crossref | Google Scholar | PubMed |

Powell JM, Morrison DA, Gadek PA, Crayn DM, Quinn CJ (1997) Relationships and generic concepts within Styphelieae (Epacridaceae). Australian Systematic Botany 10, 15-29.
| Crossref | Google Scholar |

Pritchard JK, Stephens M, Donnelly P (2000) Inference of population structure using multilocus genotype data. Genetics 155, 945-959.
| Crossref | Google Scholar | PubMed |

Puente-Lelièvre C, Hislop M, Harrington M, Brown EA, Kuzmina M, Crayn DM (2016) A five-marker molecular phylogeny of the Styphelieae (Epacridoideae, Ericaceae) supports a broad concept of Styphelia. Australian Systematic Botany 28, 368-387.
| Crossref | Google Scholar |

Radford AE, Dickison WC, Massey JR, Bell CR (1974) ‘Vascular Plant Systematics.’ (Harper & Row: New York, NY, USA)

Rouhan G, Gaudeul M (2021) Plant taxonomy: a historical perspective, current challenges, and perspectives. Methods in Molecular Biology 2222, 1-38.
| Crossref | Google Scholar | PubMed |

Rutherford S, Rossetto M, Bragg JG, McPherson H, Benson D, Bonser SP, Wilson PG (2018) Speciation in the presence of gene flow: population genomics of closely related and diverging Eucalyptus species. Heredity 121, 126-141.
| Crossref | Google Scholar | PubMed |

Scrucca L, Fop M, Murphy TB, Raftery AE (2016) mclust 5: clustering, classification and density estimation using Gaussian finite mixture models. The R Journal 8, 289-317.
| Google Scholar | PubMed |

Scrucca L, Fraley C, Murphy TB, Raftery AE (2023) ‘Model-based clustering, classification, and density estimation using mclust in R.’ (Chapman and Hall: New York, NY, USA) doi:10.1201/9781003277965

Sokal RR (1963) The principles and practice of numerical taxonomy. Taxon 12, 190-199.
| Crossref | Google Scholar |

Solbrig OT (1994) Plant traits and adaptive strategies: their role in ecosystem function. In ‘Biodiversity and Ecosystem Function.’ (Eds E-D Schulze, HA Mooney) pp. 97–116. (Springer)

Stanton DWG, Frandsen P, Waples RK, Heller R, Russo I-RM, Orozco-terWengel PA, Pedersen C-ET, Siegismund HR, Bruford MW (2019) More grist for the mill? Species delimitation in the genomic era and its implications for conservation. Conservation Genetics 20, 101-113.
| Crossref | Google Scholar |

Struck TH, Feder JL, Bendiksby M, Birkeland S, Cerca J, Gusarov VI, Kistenich S, Larsson KH, Liow LH, Nowak MD, Stedje B, Bachmann L, Dimitrov D (2018) Finding evolutionary processes hidden in cryptic species. Trends in Ecology and Evolution 33, 153-163.
| Crossref | Google Scholar | PubMed |

Taxonomy Decadal Plan Working Group (2018) ‘Discovering biodiversity: a decadal plan for taxonomy and biosystematics in Australia and New Zealand 2018–2028.’ (Australian Academy of Science: Canberra, ACT, Australia; and Royal Society Te Apārangi: Wellington, New Zealand)

von Mueller F (1868) ‘Fragmenta Phytographiae Australiae vol. VI.’ (Government Printer: Melbourne, Colony of Victoria) [In Latin] doi:10.5962/bhl.title.287

Watson L (1967) Taxonomic implications of a comparative anatomical study of Epacridaceae. New Phytologist 66, 495-504.
| Crossref | Google Scholar |

Wege JA, Thiele KR, Shepherd KA, Butcher R, Macfarlane TD, Coates DJ (2015) Strategic taxonomy in a biodiverse landscape: a novel approach to maximizing conservation outcomes for rare and poorly known flora. Biodiversity and Conservation 24, 17-32.
| Crossref | Google Scholar |