Joseph T. Miller

Molecular phylogenetics of Acacia (Fabaceae: Mimosoideae) based on the chloroplast MATK coding sequence and flanking TRNK intron spacer regions

The tribe Acacieae (Fabaceae: Mimosoideae) contains two genera, the monotypic African Faidherbia and the pantropical Acacia, which comprise about 1200 species with over 950 confined to Australia. As currently recognized, the genus Acacia is subdivided into three subgenera: subg. Acacia, subg. Aculeiferum, and the predominantly Australian subg. Phyllodineae. Morphological studies have suggested the tribe Acacieae and genus Acacia are artificial and have a close affinity to the tribe Ingeae. Based on available data there is no consensus on whether Acacia should be subdivided. Sequence analysis of the chloroplast trnK intron, including the matK coding region and flanking noncoding regions, indicate that neither the tribe Acacieae nor the genus Acacia are monophyletic. Two subgenera are monophyletic; section Filicinae of subgenus Aculeiferum does not group with taxa of the subgenus. Section Filicinae, eight Ingeae genera, and Faidherbia form a weakly supported paraphyletic grade with respect to subg. Phyllodineae. Acacia subg. Aculeiferum (s. s.) is sister to the grade. These data suggest that characters currently used to differentiate taxa at the tribal, generic, and subgeneric levels are polymorphic and homoplasious in cladistic analyses.

Phylogenetic measures of biodiversity and neo- and paleo-endemism in Australian Acacia

Understanding spatial patterns of biodiversity is critical for conservation planning, particularly given rapid habitat loss and human-induced climatic change. Diversity and endemism are typically assessed by comparing species ranges across regions. However, investigation of patterns of species diversity alone misses out on the full richness of patterns that can be inferred using a phylogenetic approach. Here, using Australian Acacia as an example, we show that the application of phylogenetic methods, particularly two new measures, relative phylogenetic diversity and relative phylogenetic endemism, greatly enhances our knowledge of biodiversity across both space and time. We found that areas of high species richness and species endemism are not necessarily areas of high phylogenetic diversity or phylogenetic endemism. We propose a new method called categorical analysis of neo- and paleo-endemism (CANAPE) that allows, for the first time, a clear, quantitative distinction between centres of neo- and paleo-endemism, useful to the conservation decision-making process.

Collapse of morphological species in the wild potato Solanum brevicaule complex (Solanaceae: sect. Petota)

The major cultivated potato, Solanum tuberosum, and six other related cultivated species, are hypothesized to have arisen from a group of weedy relatives indigenous to the central Andes of central Peru, Bolivia, and northern Argentina. A major problem hindering investigations of the origins of the cultivated species has been a continuing debate over the species boundaries of their putative progenitors. This study investigated the morphological phenetic species boundaries of these putative progenitors and five cultivated taxa, here collectively referred to as the Solanum brevicaule complex. Two hundred fifteen accessions of 30 taxa in the S. brevicaule complex and 42 accessions of six taxa outside of the complex were assessed for 53 morphological traits in replicate plots in a common garden, resulting in a total of over 81;t3000 data points. Phenetic analyses of these data are unable to support 30 taxa, suggesting instead a single variable complex at best only weakly divided into three widely intergrading sets of populations: (1) Peruvian and geographically adjacent Bolivian accessions (including wild species and all the cultigens), (2) Bolivian and Argentinian accessions and S. verrucosum from Mexico (including only wild species), and (3) the Bolivian and Argentinian wild species S. oplocense. These and other data suggest that Hawkess 1990 treatment (The Potato: Evolution, Biodiversity, and Genetic Resources, Smithsonian Institute Press, Washington, DC.) of 232 morphological species is an overestimate for sect. Petota.

Acacia s.s. and its Relationship Among Tropical Legumes, Tribe Ingeae (Leguminosae: Mimosoideae)

Abstract To search for the sister taxon of Acacia s.s. (tribe Acacieae) and to further knowledge of the phylogeny of the related tribe Ingeae, we have sequenced two regions of nuclear ribosomal DNA (ITS and ETS). Sixty species from tribe Ingeae (26 genera), together with representatives from each of five lineages of tribe Acacieae, have been sampled. Ingeae and Acacia s.s. form a well supported clade, with a monophyletic Acacia s.s. nested within a paraphyletic Ingeae. Based on our sampling, the closest relative of Acacia s.s. is most likely one of the Australian species of the genus Paraserianthes s.l.: Paras. lophantha subsp. lophantha or Paras. toona. Related to Acacia s.s. and Paraserianthes s.l. is a group of Ingeae from Australia and South East Asia: Archidendron p.p., Archidendropsis, Pararchidendron, and Wallaceodendron. This study is a preliminary step in resolving the intergeneric relationships of tribe Ingeae. Genetic relationships within Ingeae appear to conform to morphological groups previously identified as genera and informal alliances; some notable exceptions are discussed.

Molecular phylogeny of Acacia subgenus Phyllodineae (Mimosoideae : Leguminosae) based on DNA sequences of the internal transcribed spacer region

The largest monophyletic group within Acacia is subgenus Phyllodineae, with more than 950 predominately Australian species, the majority characterised by adult foliage consisting of phyllodes. Molecular sequence data from the internal transcribed spacers (ITS) of the nuclear ribosomal DNA repeat were used to investigate the monophyly of seven sections within the subgenus. A nested PCR approach was used to amplify the ITS region. Fifty-one species representative of all sections were sequenced together with one outgroup taxon Lysiloma divaricata (Ingeae). Phylogenetic parsimony analysis suggested that there are two main clades within Phyllodineae but that only one section, Lycopodiifoliae, is apparently monophyletic. In one of the main clades, Lycopodifoliae is related to some taxa in sections Alatae and Pulchellae and some members of section Phyllodineae. In the second main clade, sections Juliflorae, Plurinerves and Botrycephalae cluster with other members of section Phyllodineae. The two sections that are characterised by bipinnate foliage, Botrycephalae and Pulchellae, are nested within phyllodinous clades, indicating that at least two separate reversals to bipinnate leaves have occurred. Botrycephalae is paraphyletic with respect to taxa from section Phyllodineae that have single-nerved phyllodes and racemose inflorescences.

Spatial distribution of species richness and endemism of the genus Acacia in Australia

The aim of this study is to identify and map the spatial distribution of species richness and endemism of the genus Acacia in Australia. A database of 171 758 geo-referenced herbarium records representing 1020 Acacia species was assembled and aggregated to a 0.25° grid cell resolution. A neighbourhood analysis of one-cell radius was applied to each of the grid cells to map the spatial patterns of species richness and endemism. The primary centres of species richness are in accordance with previous results, occurring in the South-West Botanical Province in Western Australia, the MacPherson-Macleay overlap and the Central Coast of the Sydney Sandstone region. We identify 21 centres of endemism, of which six were previously unrecognised. The primary centres of endemism are located in South-West Western Australia, the Kimberley District and the Wet Tropics in Queensland. The South-West Botanical Province in Western Australia contained the greatest number of regions with the highest number of endemic species of Acacia. A randomisation test showed that our 21 centres of endemism were significantly different from random. The majority of centres of Acacia endemism were incongruent with the centres of species richness, with only three grid cells in the top 1% for both measures. We also confirm that South-West Western Australia is a region of very high species richness and endemism, in accordance with its status as a global hotspot of biodiversity.

Molecular phylogenetics of Acacia subgenera Acacia and Aculeiferum (Fabaceae : Mimosoideae), based on the chloroplast matK coding sequence and flanking trnK intron spacer regions

The genus Acacia is subdivided into the following three subgenera: subg. Acacia, subg. Aculeiferum and the predominantly Australian subg. Phyllodineae. Morphological and molecular studies have suggested that the tribe Acacieae and genus Acacia are artificial and have a close affinity to the tribe Ingeae. Sequence analysis of the chloroplast trnK intron, including the matK coding region and flanking non-coding regions, were undertaken to examine taxon relationships within Acacia subgenera Acacia and Aculeiferum. Subgenus Acacia is monophyletic while subgenus Aculeiferum is paraphyletic. Within the subgenera, major divisions are found based on biogeography, New World versus African-Asian taxa. These data suggest that characters such as inflorescence and prickle and/or stipule type are polymorphic and homoplasious in cladistic analyses within the subgenera. SB lo f Jd

Phylodiversity to inform conservation policy: An Australian example

Phylodiversity measures summarise the phylogenetic diversity patterns of groups of organisms. By using branches of the tree of life, rather than its tips (e.g., species), phylodiversity measures provide important additional information about biodiversity that can improve conservation policy and outcomes. As a biodiverse nation with a strong legislative and policy framework, Australia provides an opportunity to use phylogenetic information to inform conservation decision-making. We explored the application of phylodiversity measures across Australia with a focus on two highly biodiverse regions, the south west of Western Australia (SWWA) and the South East Queensland bioregion (SEQ). We analysed seven diverse groups of organisms spanning five separate phyla on the evolutionary tree of life, the plant genera Acacia and Daviesia, mammals, hylid frogs, myobatrachid frogs, passerine birds, and camaenid land snails. We measured species richness, weighted species endemism (WE) and two phylodiversity measures, phylogenetic diversity (PD) and phylogenetic endemism (PE), as well as their respective complementarity scores (a measure of gains and losses) at 20 km resolution. Higher PD was identified within SEQ for all fauna groups, whereas more PD was found in SWWA for both plant groups. PD and PD complementarity were strongly correlated with species richness and species complementarity for most groups but less so for plants. PD and PE were found to complement traditional species-based measures for all groups studied: PD and PE follow similar spatial patterns to richness and WE, but highlighted different areas that would not be identified by conventional species-based biodiversity analyses alone. The application of phylodiversity measures, particularly the novel weighted complementary measures considered here, in conservation can enhance protection of the evolutionary history that contributes to present day biodiversity values of areas. Phylogenetic measures in conservation can include important elements of biodiversity in conservation planning, such as evolutionary potential and feature diversity that will improve decision-making and lead to better biodiversity conservation outcomes.

Phylogenetic diversity meets conservation policy: Small areas are key to preserving eucalypt lineages

Evolutionary and genetic knowledge is increasingly being valued in conservation theory, but is rarely considered in conservation planning and policy. Here, we integrate phylogenetic diversity (PD) with spatial reserve prioritization to evaluate how well the existing reserve system in Victoria, Australia captures the evolutionary lineages of eucalypts, which dominate forest canopies across the state. Forty-three per cent of remaining native woody vegetation in Victoria is located in protected areas (mostly national parks) representing 48% of the extant PD found in the state. A modest expansion in protected areas of 5% (less than 1% of the state area) would increase protected PD by 33% over current levels. In a recent policy change, portions of the national parks were opened for development. These tourism development zones hold over half the PD found in national parks with some species and clades falling entirely outside of protected zones within the national parks. This approach of using PD in spatial prioritization could be extended to any clade or area that has spatial and phylogenetic data. Our results demonstrate the relevance of PD to regional conservation policy by highlighting that small but strategically located areas disproportionally impact the preservation of evolutionary lineages.

Genetic, cytogenetic and morphological patterns in a mixed mulga population: evidence for apomixis

The mulga complex (Acacia aneura and closely related taxa) is a widespread group that is dominant in much of arid Australia. The group is taxonomically difficult, due to a complex interaction of sympatry and putative hybridisation between the major species, geographic variation within species and sympatric variation within A. aneura. Mulga is highly variable in a wide range of vegetative and reproductive characters and it is not unusual to find five or six distinct forms growing side by side. The aim of this project was to gain a better understanding of the relationships among mulga species and A. aneura varieties, as well as the maintenance of this variation. A single site in the Northern Territory, containing A. ayersiana, A. minyura and two varieties of A. aneura, was sampled intensively. Six morphotypes were observed in the field and five were strongly supported by morphometric analysis. Although the mulga complex is generally tetraploid (2n = 52), triploid (2n = 39) and pentaploid (2n = 65) seedlings were produced in the study population. Microsatellite primers developed for A. mangium (sect. Juliflorae) were amplified in individuals of each morphotype, resulting in genetic marker patterns consistent with polyploidy. Genetic and morphometric distances were correlated and differences between morphotypes account for 63% of the total genetic variation (φPT = 0.63, P < 0.001). Allele sequences confirmed the presence of genuine heterozygosity and clonality was suggested by the low genotypic diversity and the lack of allele segregation. Seedlings had identical genotypes to the maternal plants and polyembryony was observed in each taxon, consistent with apomictic reproduction. Both apomixis and ploidy level variation may restrict gene flow among morphotypes, playing a role in the maintenance of morphological diversity at the study site. The success of the group in arid and semi-arid Australia may also be due, in part, to these factors. SB enet or v ul R. L.