Corresponding author: Gillian K. Brown (
Academic editor: Colin E. Hughes
The morphologically variable genus
Brown GK, Aju J, Bayly MJ, Murphy DJ, McLay TGB (2022) Phylogeny and classification of the Australasian and Indomalayan mimosoid legumes
The classification of mimosoid legumes has been significantly transformed in the past 20 years since the first comprehensive molecular phylogeny of the then subfamily
The two largest mimosoid genera from the Indomalayan-Australasian region are
Distribution maps of the genera
Prior to resolution of the
The genus
This study aims to test the monophyly of the genera
A total of 87 accessions were sampled, representing 43 species of
Linked data table of specimens sampled for phylogeny. Specimen accession number linking herbarium specimen to sample ID, taxon name with authorities, locality information and geocode (where available) as provided on the specimen/database. GenBank numbers are provided for each marker and where multiple alleles were identified for a specimen, the two GenBank numbers are separated by a semi colon. If the marker was not successfully sequenced for a particular specimen, then the GenBank field is left blank.
Preserved specimen | Associated sequences | Taxon name/MOTU | Sample ID | Location | |||||||
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Specimen code (InstCode and/or CollCode + Catalogue #) | SHMT | RBPCO | ITS | ETS | trnK | trnV | psbD | Geolocation name / locality | GPS Coordinates | ||
MEL 2294706A |
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Great Sandy National Park, Fraser Island, Woralie track to Moon Point. Queensland, Australia |
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MELU GB309b |
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0.7km north of Playford Highway on Snug Bay Rd, Kangaroo Island, South Australia |
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CANB 864530.1 |
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Alva, NE of Ayr, Queensland, Australia |
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MEL 2391890A |
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Atherton Arboretum. Tag #96. Queensland, Australia |
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KUN0599506 |
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China |
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BRI AQ0380081 |
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Papua New Guinea, Western Fly; Kwinja Lakes area of the Middle Fly River |
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KUN0599551 |
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China | |||||
AAU D.McKey92-9 |
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Sinharaja Forest, SW Sri Lanka |
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AAU Balgooy6063 |
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Tanah Merah, Kalimantan Timur |
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KEP FRI53789 |
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Pahang, Temerloh, Tasik Bera, Kg. Patihir, Malaysia |
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CANB 730419.1 |
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Ambunti District, Waskut Hills, spur ridge NW of Musapien bivouac. East Sepik, PNG |
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CANB 211609.1 |
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Saw Mountains, near junction of Tauri and Kapau Rivers. Gulf Province, PNG |
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AAU L.Averyanov4481 |
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Bi Dup ridge, Vietnam |
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AAU I.Nielsen26 |
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Gunung Mulu National Park, Sarawak |
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AAU H.M.Christensen38 |
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Pa Dalih area, Sarawak |
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AAU L.AveryanovVH3188 |
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Bi Dup mountain system, Vietnam |
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CANB 525617.1 |
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East branch of the Avi Avi River. Gulf Province, PNG |
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AAU AmbriW838 |
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Wanariset research area, Kalimantan Timur |
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NY03986843 |
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Near Kuala Lumpur, Malaysia | |||
BRI AQ0380332 |
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Papua New Guinea, Central; Subitana, Sogeri sub-dist., Central, Papua |
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BISH752370 |
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Siboma, Sayama, track along the ridgeline S from Camp 1. PNG |
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BISH763497 |
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Morobe Province; Oomsis, behind PNG Forestry station. |
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BRI AQ0380375 |
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Brown River F.R. Central Province, PNG |
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AAU J.F.Maxwell82-141 |
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Near Bukit Kallang, Singapore | |||
AAU Bjornland445 |
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Chiang Mai: Amphoe Muang, Mae Rim, Thailand | ||||
CANB 544379.1 |
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Gabba Island, Torres Strait. Queensland, Australia |
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BRI AQ0814833 |
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Curramore Sanctuary Nature Reserve, 14km NW of Maleny. Queensland, Australia |
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CNS 131336.1 |
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Mt Lewis, Carbine Tableland. Queensland, Australia |
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MEL 2391892A |
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Atherton Arboretum. Tag #846. Queensland, Australia |
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AAU Kostermans22121 |
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Mbengen, West Flores | |||||
AAU H.M.Christensen279 |
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Pa Dalih area, Sarawak |
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CANB 596487.1 |
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Greenfield Road, Lennox Head. New South Wales, Australia |
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MEL 2293327A |
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Brandy Creek Road, 9 km S of Airlie Beach. Queensland, Australia |
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QRS 18805.2 |
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Between Starcke homestead and Starcke River. Queensland, Australia |
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MEL 2391969A |
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Cairns, cultivated in garden. Queensland, Australia |
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QRS 117169.1 |
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Claudie River. Queensland, Australia |
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CNS 142441.1 |
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Umagico, Cape York. Queensland, Australia |
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MEL 2391887A |
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Atherton Arboretum. Tag #482. Queensland, Australia |
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BISH760310 |
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Northern Province; Sibium Mountains; W of Akupe Camp, along Afase River. PNG |
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AAU R.Geesink7254 |
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Kao Chong Botanical Garden, Thailand |
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BRI AQ0738090 |
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Oliver Creek. Queensland, Australia |
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MELUD113392a |
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Shore of creek, end of Stonewood Road, Queensland, Australia |
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MELUD113385a |
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Shore of creek, end of Stonewood Road, Queensland, Australia |
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BRI AQ0733240 |
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NPR133, Daintree, Oliver Creek, Queensland, Australia |
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AAU I.Cowley110 |
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Melilas. Ulu Belait, Brunei | ||||
AAU H.M.Christensen1719 |
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near Nanga Sumpa, Sarawak |
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KUN 0599659 |
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BRI AQ0835639 |
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Great Sandy National Park; Cooloola Section, Freshwater Road. Queensland, Australia. |
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BRI AQ0636343 |
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Harry’s Hut Road, Cooloola National Park.Queensland, Australia |
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MEL 2034578A |
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Indooroopilly, cultivated. Queensland, Australia | |||
MELUD113387a |
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Lake Road near Cairns, Queensland, Australia |
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MELUD113393a |
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Lake Road near Cairns, Queensland, Australia |
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MELUD113391a |
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Cape Tribulation Road, adjacent to Coconut Beach resort, Queensland, Australia |
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MEL 2391968A |
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Cairns, cultivated in garden. Queensland, Australia |
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BISH760584 |
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Central Province; Mt Gerebu, trail towards summit ridge. PNG |
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AAU H.M.Christensen1282 |
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Near Sumpa, Sarawak. |
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BRI AQ0499073 |
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Big Scrub Flora Reserve, NNE of Lismore, New South Wales, Australia |
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BRI AQ0763292 |
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Tallebudgera Creek Road, reveg site. Queensland, Australia |
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BISH752405 |
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Morobe Province; Siboma, Sayama, above Sayama Creek, to E Camp 1. PNG |
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MEL 2074350A |
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N. de Dalat, prov. Ht. Donnai. Indochina: Annam. Vietnam |
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Bell Museum 913425 (WP-3A0575) |
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Wanang village, Madang, PNG |
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AAU C.Charoenphol5025 |
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Ko Rang Yai, Thailand |
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MEL 2391884A |
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Atherton Arboretum. Tag #1652. Queensland, Australia |
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MELUD113388a |
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Regeneration plot, Daintree Rainforest Observatory, Queensland, Australia |
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BRI AQ0485087 |
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Shiptons Flat. Queensland, Australia |
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AAU Balgooy6769 |
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Pulan Baun, Aru Island Indonesia |
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BRI AQ0052837 |
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Agu River branch of the middle Fly River, PNG |
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MEL 2041191A |
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Cape Tribulation, Queensland, Australia |
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BRI AQ0558405 |
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Along Paluma Dam Road, Ethel Creek, Queensland, Australia |
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MEL 2196304A |
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State Forest 310 Gadgarra. Queensland, Australia |
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BRI AQ0824396 |
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7km W of Babinda, Queensland, Australia. |
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KUN 0599686 |
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China | ||||
BRI AQ0611431 |
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On Isaac River and Hill Creek. 25km S of Glenden, Queensland, Australia |
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MEL 0290000A |
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Bladensburg National Park, S of Winton, Poison Paddock. Queensland, Australia |
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MEL 2333247A |
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Prov. Sud, near Yate, north side of Yate River. New Caledonia |
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BRI AQ0430532 |
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c. 5km north of Kone, south of Kafeate. New Caledonia. |
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MEL 2095888A |
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Palmgrove National Park, 5km W of Daydream Hill, Queensland, Australia |
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BRI AQ0771148 |
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Daintree, narrow ridge above Cassowary Creek, off Stewart Creek road, site 69. Queensland, Australia |
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L.1958248 |
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KPC area, Sebongkok Utara, East Kalimantan, Indonesia. |
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CANB 367091.1 |
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Near Earlando, 27 km N of Proserpine. Queensland, Australia |
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MEL 1615244A |
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Palm Tree Creek, W of Mt Whitestone township, Queensland, Australia |
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CNS 134531.1 |
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CSIRO Arboretum, Queensland, Australia |
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QRS 121813.1 |
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Clarke Range, Queensland, Australia |
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MEL 2183015A |
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Merrimu Reservoir, Victoria, Australia |
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BRI AQ0408829 |
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Atop Sailigai Hulo, Rota. Northern Mariana Islands. |
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MEL 2333248A |
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Prov. Sud, near Prony, New Caledonia |
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MELU SRA051 |
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Bogor Botanic Gardens collection Accession: |
Total genomic DNA (
Eight nuclear markers (low copy genes: AIGP, CYB6, Eif3E, SHMT, RBPCO, UDPG; nrDNA: ITS, ETS) and four chloroplast DNA intergenic spacer regions (
PCR reagents, primers and cycling conditions are described in Suppl. material
Library preparation followed the two-step PCR process outlined in
The second step used qPCR to add unique Illumina indexing barcodes to each sample for the pooled amplicons. Indexing PCR reactions consisted of 5 µM of each of index primer (
Sequences obtained by Sanger sequencing were aligned by individual locus in Geneious v.8.1.4 (Biomatters Ltd.) and a consensus sequence was generated and used as the reference for the reads obtained by
Alignments of all nuclear loci (
A consensus network of the combined
Partition | Model |
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ETS | HKY+F+G4 |
ITS1 | GTR+F+G4 |
5.8S | SYM+I+G4 |
ITS2 | HKY+F+G4 |
SHMT | HKY+F+G4 |
RBPCO | K2P+I |
All chloroplast (
Pollen size and surface texture are key morphological features differentiating the subgenera of
Of the eight nuclear loci only four were included in the final phylogenetic analyses: SHMT, RBPCO, ITS and ETS. ETS and ITS amplified well, were variable, and are commonly used phylogenetic markers in
Of the four chloroplast loci,
The topologies of the combined
Combined
Within
Of the 12 species of
The consensus network of the final 101-sampled
The phylogeny of the three
Combined
The pollen measurement results are consistent with
Scanning electron micrographs of
Our study presents the most taxon-rich sampling of the
Despite the poorly resolved backbone of the
The genus
These three subclades of the
Three species in the
The
The last revision of the genus
While
It was only recently that the species of the two subgenera were united within
The presence of a pleurogram is common in mimosoid genera (
The non-monophyly and clear morphological distinctions between them means that the two subgenera can no longer be treated as congeneric and need to be placed in separate genera. As the type of
In addition to the aforementioned morphological differences between the two subgenera, species of subg.
Morphology of
Flowers arranged in globular heads, seeds lacking a pleurogram with a narrow peripheral membranous wing and flat, narrowly oblong, brown pods opening along both sutures distinguish this new genus from other Australian mimosoid legumes, and the keys in Flora of Australia (
A genus of mimosoid legumes similar to
Trees or shrubs, with terete branchlets. Stipules either resembling small thorns to 1.2 mm long that are early caducous, or persistent circular-ovate glands 1–3 mm in diameter. Leaves bipinnate, pinnae 1–2 pairs with 1.5–11 leaflet pairs per pinna; glands at the junction of pinnae circular or triangular to rhombic, +/- circular glands at the junction of leaflet petiolules. Leaflets opposite, subsessile (0.2–0.7 mm) or long (3.5–7 mm) petiolulate; elliptic to elliptic-lanceolate or oblong, 2–38 mm × 1.5–15 mm, glabrous to puberulous. Inflorescence of globular heads 0.5–1.7 mm in diameter, either simple or arranged into a panicle up to 35 cm long. Flowers: homomorphic, yellow to cream, sessile. Calyx 1.5–3 mm long, tubular to subcampanulate; corolla 2.5–7 mm long, tubular to narrowly campanulate. Ovary 0.8–2 mm long, solitary and shortly stipitate; stamens numerous 5–9 mm long, united basally into a tube that equals or slightly exceeds the corolla tube. Pollen 16-celled polyads with a diameter of 55–68 μm, tectum with isometric channels. Pod brown, valves chartaceous, 6–22 cm × 0.5–2.5 mm, oblong, flat and dehiscing along both sutures. Seeds lacking a pleurogram, flat, circular to ovate or obliquely ovate, 5–13 mm, with a narrow 0.2–1 mm peripheral, membranous wing. Fig.
From the Greek
We have chosen to create a new name for this genus rather than making a new combination based on the name
The genus includes the following three species, all endemic to Queensland, Australia (Fig.
Peak Downs,
Fort Cooper, [
Atherton District, North Queensland,
The protologue of
In Flora of Australia,
We present the most densely sampled phylogeny of the genera
This research was funded by an Australia and Pacific Science Foundation grant (APSF14-4: Systematics and evolution of
Primer sequences and PCR variations
Pdf file.
The reference for the primer and their PCR conditions are provided, along with the variations for PCR reagents and cycling conditions for the initial PCR in the two-step PCR process. * only used for sanger sequencing so no variations to note. Standard PCR reagents prior to variation consisted of 2X QIAGEN PCR buffer (QIAGEN), 5 mM of each dNTP (Bioline), 1 µl of each primer (10 µM), 1.25 µl of dimethyl sulfoxide (DMSO, 99.5%; Sigma-Aldrich), 1 U of Taq DNA polymerase, 100 ng of template and made up to 25 µl with ultra pure water per reaction. Reagent variations, A: not varied; B: 200 ng DNA, 1.2 µl BSA instead of DMSO; C: 200 ng DNA; D: 200ng DNA, 6 μM each primer, 1.5 µl MgCl2, 0.9 µl DMSO and 0.1 µl Taq; E: 6 µM each primer. Cycle variations: Z: 94 °C for 15 mins; 30 cycles of 94 °C for 20 sec, 61 °C for 20 sec, 72 °C for 2 mins; 72 °C for 5 mins; Y: 94 °C for 15 mins; 35 cycles of 94 °C for 20 sec, 61 °C for 20 sec, 72 °C for 2 mins; 72 °C for 5 mins; X: 94 °C for 15 mins; 35 cycles of 94 °C for 20 sec, 55 °C for 30 sec, 72 °C for 2 mins; 72 °C for 7 mins; W: 94 °C for 15 mins; 40 cycles of 94 °C for 20 sec, 50 °C for 1 min, 72 °C for 3 mins; 72 °C for 7 mins; V: 80 °C for 5 mins; 40 cycles of 95 °C for 1 min, 50 °C for 1 min with 0.3 °C/sec ramp, 65 °C for 4 mins; 65 °C for 5 mins; U: 94 °C for 5 mins; 30 cycles of 94 °C for 30 sec, 53 °C for 30 sec, 72 °C for 1 min; 72 °C for 7 mins; T: 80 °C for 5 mins; 30 cycles of 95 °C for 1 min, 50 °C for wwith 0.3 °C/sec ramp, 65 °C for 4 mins; 65 °C for 5 mins.
SHMT network and tree
Pdf file.
Neighbour-joining tree and NeighbourNet network are presented with individual samples with more than one allele coloured to highlight their positions. The samples are coloured the same in both the tree and network. The clades that are congruent with Fig.
RBPCO network and tree
Pdf file.
Neighbour-joining tree and NeighbourNet network are presented with individual samples with more than one allele coloured to highlight their positions. The samples are coloured are the same in both the tree and network. Clade B, which is congruent with Fig.
Pdf file.
IQ-Tree of combined