Pauline Y. Ladiges

Stemming from acceptance of continental drift theory and development of cladistic systematics in the 1960s, interest in historical biogeography was revived. The field and methodology have since been reviewed several times (e.g., Nelson and Platnick 1981; Myers and Giller 1988; Morrone and Carpenter 1994; Humphries and Parenti 1999; Crisci et al. 2003; Ebach and Tangney 2007; Santos 2007; Parenti and Ebach 2009). In essence, early methods focussed on discovering relationships of areas based on phylogenies of taxa endemic to those areas and biogeographic patterns presented as summary area cladograms. Comparisons across different groups (clades) of organisms revealing congruent patterns were interpreted as evidence of simple vicariance, which could be compared with other data, such as geological breakup of Gondwana, whereas incongruence of patterns was interpreted as requiring a complex explanation or stochastic long-distance dispersal (Nelson and Platnick 1981). A plethora of methods to determine area cladograms arose to deal with widespread taxa (representing range expansions or a failure to differentiate), missing areas (extinctions), and other complications. But by the 1990s, there was no unifying accepted biogeographic method (Nelson and Ladiges 2001), and different methods applied to the same data gave different results, indicating that at least some of the results are artifacts of the method.

Published phylogenies of two eucalypt clades, red bloodwoods Corymbia subgenus Corymbia and eudesmids Eucalyptus subgenus Eudesmia (Myrtaceae), were combined for an analysis of historical biogeographical area relationships within continental Australia. The method of paralogy-free subtree analysis was used to eliminate geographical paralogy; the paralogy-free subtrees were coded as characters for parsimony analysis to find the minimal and area cladogram, which proved to be informative of a continent-wide pattern. The eucalypt fossil record and molecular dating studies allow an interpretation of the biogeographical history in terms of major vicariance events that date from the early Paleogene. The summary area cladogram shows the wet jarrah forest region of South-West Western Australia, a region of high endemism, as the earliest to differentiate from all other areas, isolated by marine inundation across southern Australia and climatic cooling in the Late Eocene-Early Oligocene. From about this time, regionalization continued, with warmer conditions and monsoonal climate developing in central and northern Australia, and cooling in the south-east. Northern and eastern humid and semi-humid areas were related as a track, but with increased aridity in the interior of the continent, the monsoonal climate contracted northwards. The Australian Monsoon Tropics (AMT: Kimberley, Top End, Arnhem, Cape York and inland north-east Queensland) differentiated from eastern areas (Queensland wet tropics to McPherson-Macleay). Our results also show all arid and semi-arid regions as related, suggestive of a historically cohesive interior biota rather than repeated colonizations of the interior from the periphery of the continent. Climate largely differentiates hot arid areas in the north (Pilbara, Northern and Central deserts) from arid areas in the south (south-west interzone, Wheatbelt, Goldfields and Great Victoria Desert). (C) The Willi Hennig Society 2010.

Phylogenetic analysis of Eucalyptus subgenus Eudesmia is presented on the basis of the following three datasets: sequences of the internal transcribed spacer (ITS) and the external transcribed spacer (ETS) regions from nuclear rDNA, sequences of the psbA–trnH intergenic spacer region from chloroplast DNA, and morphological characters, including stamen bundling, operculum development, seeds and trichomes. Studies of floral development were essential for understanding the morphology of mature flowers and interpretation of synapomorphy and homoplasy. A summary phylogeny was constructed from a maximum parsimony analysis of those nodes coded as characters that had support in the molecular trees together with morphological characters. A revised infra-subgeneric classification is presented on the basis of the summary phylogeny, and compared with classifications of Hill and Johnson (1998) and Brooker (2000). Differences relate to relationships between clades and taxonomic rank (sections, series and subseries) and valid names of Brooker (2000) are conserved where possible. One main clade of 14 species (section Limbatae), many of mallee growth form, was found in all analyses; this clade is distributed in the South-West of Western Australia and adjacent Interzone and desert areas. A second main clade (section Complanatae) occurs in the northern and eastern tropical and subtropical regions of Australia, including Kimberley, Arnhem, Queensland and New South Wales. This section includes E. tetrodonta, previously treated as an isolated taxon in a monotypic section; however, this species is related to E. baileyana, E. similis, E. lirata and series Miniatae. The hypothesised phylogeny provides a framework for further analyses of biogeography and ecology, including functional traits.

Phylogenetic relationships of sections and species within Corymbia (Myrtaceae), the bloodwood eucalypts, were evaluated by using combined analyses of nuclear rDNA (ETS + ITS) and morphological characters. Combining morphological characters with molecular data provided resolution of relationships within Corymbia. The analyses supported the monophyly of the genus and recognition of the following two major clades, treated here as new subgenera: subgenus Corymbia, including informal sections recognised by Hill and Johnson (1995), namely Rufaria (red bloodwoods), Apteria and Fundoria; and subgenus Blakella, including sections Politaria (spotted gums), Cadagaria, Blakearia (paper-fruited bloodwoods or ghost gums) and Ochraria (yellow bloodwoods). Hill and Johnson’s section Rufaria is monophyletic if Apteria and Fundoria are included. It is evident that, among the red bloodwoods, series are not monophyletic and several morphological characters result from convergent evolution. There was strong morphological and molecular evidence that the three species of red bloodwoods that occur in south-western Western Australia (series Gummiferae: C. calophylla and C. haematoxylon, and series Ficifoliae: C. ficifolia) form a monophyletic group, separate from the eastern C. gummifera (series Gummiferae), which is probably sister to the clade of all other red bloodwoods. Phylogenetic results supported recognition of new taxonomic categories within Corymbia, and these are formalised here.

A review is presented of current knowledge of the tectonic rifting of eastern Gondwana in the context of possible land continuity between Australia and New Caledonia and the dating of events. Comment is made on the level of endemicity of the modern-day biota of New Caledonia, with examples of phylogenetic analyses, biogeographic patterns and estimated divergence dates among particular taxa, including birds (kagu-sunbittern clade), Nothofagus and Myrtaceae. The geological history of the region is complex but there is evidence of land persisting into the Cenozoic (Paleocene/Eocene) that possibly allowed old biota to persist. Vicariance explanations of divergences cannot be justifiably dismissed in favour of long-distance, over-water dispersal for such taxa.

Morphological variation in Acacia victoriae, a species widespread in arid and semi-arid regions of Australia, was analysed by phenetic methods of classification and ordination. Three morphological groups were identified on the basis of phyllode characters and are treated as subspecies. Populations with short, elliptic and tomentose phyllodes are confirmed as A. victoriae subsp. arida Pedley; this form occurs mainly in central Australia. Populations with linear to oblong, non-tomentose phyllodes are referred to subsp. victoriae; this subspecies is the most variable and widely distributed across Australia. Populations with very long, narrow phyllodes, distributed in northern Australia from the Kimberley to Queensland, are described as fasciaria subsp. nov. A small number of non-tomentose specimens with broad elliptic phyllodes from central Australia require further assessment. Thirteen accessions previously sequenced for internal spacer regions and external spacer regions of ribosomal nuclear DNA showed genetic divergence. Six accessions of subsp. fasciaria formed a clade in a parsimony analysis, confirming that the long phyllode form is genetically distinct.

The phylogeny of three groups of arid Australian acacias ‐ the Acacia victoriae, A. murrayana and A. pyrifolia groups ‐ was constructed based on parsimony analysis of sequence data from the internal and external transcribed spacers (ITS and ETS) of the nuclear ribosomal DNA. Forty ingroup taxa were sequenced, including multiple accessions for some taxa and two species (A platycarpa and A. longispinea) that had been identified in other analyses as relatives of these acacias. Acacia anthochaera was used as the functional outgroup. The ITS and ETS regions proved to be sufficiently variable to resolve relationships at both the specific and intra‐specific level. Two main clades were resolved. One clade confirmed the monophyly of the Acacia murrayana group, and relationships of species were strongly supported. All taxa in this clade have a similar pattern of seedling leaf development. In the second clade, the A. pyrifolia group is nested within the A. victoriae group and all taxa have spinose stipules. Acacia platycarpa and A. longispinea are related to this clade. Phyllode nerve number (uninerved or plurinerved) proved to be homoplasious. Acacia victoriae is a widespread and very variable species. The molecular data identified two major groups: a group of populations occurring across northern Australia and a group of populations from the Western, Central and Eastern deserts. Further analysis of population variation is required to assess the taxonomic status of various forms in this species complex. The geographic distributions of sister taxa suggest predominantly allopatric speciation. The degree of molecular divergence and position of the clades within subgenus Phyllodineae suggest that the lineages are not of recent origin, but have a history that relates to increased aridity in the Australian Eremean region during the Cenozoic.