Process and pattern in the biogeography of New Zealand – a global microcosm?

Aim  To describe New Zealand's historical terrestrial biogeography and place this history in a wider Southern Hemisphere context.
Location  New Zealand.
Methods  The analysis is based primarily on literature on the distributions and relationships of New Zealand's terrestrial flora and fauna.
Results  New Zealand is shown to have a biota that has broad relationships, primarily around the cool Southern Hemisphere, as well as with New Caledonia to the north. There are hints of ancient Gondwanan taxa, although the long-argued predominance of taxa derived by vicariant processes, driven by plate tectonics and the fragmentation of Gondwana, is no longer accepted as a principal explanation of the biota's origins and relationships.
Main conclusions  Most of the terrestrial New Zealand flora and fauna has clearly arrived in New Zealand much more recently than the postulated separation of New Zealand from Gondwana, dated at c. 80 Ma. There is a view that New Zealand may have disappeared completely beneath the sea in the early Cenozoic, and acceptance of this would mean derivation of the entire biota by transoceanic dispersal. However, there are elements in the biota that seem to have broad distributions that date back to Gondwanan times, and also some that are thought unlikely to have been able to disperse to New Zealand across ocean gaps, especially freshwater organisms. Very strong connections to the biota of Australia, rather than to South America, are inconsistent with the timing of New Zealand's ancient and early separation from Gondwana and seem likely to have resulted from dispersal.     

Biogeography of the Monimiaceae (Laurales): a role for East Gondwana and long‐distance dispersal,but not West Gondwana

Aim The biogeography of the tropical plant family Monimiaceae has long been thought to reflect the break‐up of West and East Gondwana, followed by limited transoceanic dispersal. Location Southern Hemisphere, with fossils in East and West Gondwana. Methods We use phylogenetic analysis of DNA sequences from 67 of the c. 200 species, representing 26 of the 28 genera of Monimiaceae, and a Bayesian relaxed clock model with fossil prior constraints to estimate species relationships and divergence times. Likelihood optimization is used to infer switches between biogeographical regions on the highest likelihood tree. Results Peumus from Chile, Monimia from the Mascarenes and Palmeria from eastern Australia/New Guinea form a clade that is sister to all other Monimiaceae. The next‐deepest split is between the Sri Lankan Hortonia and the remaining genera. The African Monimiaceae, Xymalos monospora, then forms the sister clade to a polytomy of five clades: (I) Mollinedia and allies from South America; (II) Tambourissa and allies from Madagascar and the Mascarenes; (III) Hedycarya, Kibariopsis and Leviera from New Zealand, New Caledonia and Australia; (IV) Wilkiea, Kibara, Kairoa; and (V) Steganthera and allies, all from tropical Australasia. Main conclusions Tree topology, fossils, inferred divergence times and ances‐tral area reconstruction fit with the break‐up of East Gondwana having left a still discernible signature consisting of sister clades in Chile and Australia. There is no support for previous hypotheses that the break‐up of West Gondwana (Africa/South America) explains disjunctions in the Monimiaceae. The South American Mollinedia clade is only 28–16 Myr old, and appears to have arrived via trans‐Pacific dispersal from Australasia. The clade apparently spread in southern South America prior to the Andean orogeny, fitting with its first‐diverging lineage (Hennecartia) having a southern‐temperate range. The crown ages of the other major clades (II–V) range from 20 to 29 Ma, implying over‐water dispersal between Australia, New Caledonia, New Zealand, and across the Indian Ocean to Madagascar and the Mascarenes. The endemic genus Monimia on the Mascarenes provides an interesting example of an island lineage being much older than the islands on which it presently occurs.     

Age and historical biogeography of the pantropically distributed Spathelioideae (Rutaceae,Sapindales)

Aim The family Rutaceae (rue family) is the largest within the eudicot order Sapindales and is distributed mainly in the tropical and subtropical regions of both the New World and the Old World, with a few genera in temperate zones. The main objective of this study is to present molecular dating and biogeographical analyses of the subfamily Spathelioideae, the earliest branching clade (which includes eight extant genera), to interpret the temporal and spatial origins of this group, ascertaining possible vicariant patterns and dispersal routes and inferring diversification rates through time. Location Pantropics. Methods A dataset comprising a complete taxon sampling at generic level (83.3% at species level) of Spathelioideae was used for a Bayesian molecular dating analysis (beast ). Four fossil calibration points and an age constraint for Sapindales were applied. An ancestral area reconstruction analysis utilizing the dispersal–extinction–cladogenesis model and diversification rate analyses was conducted. Results Dating analyses indicate that Rutaceae and Spathelioideae are probably of Late Cretaceous origin, after which Spathelioideae split into a Neotropical and a Palaeotropical lineage. The Palaeotropical taxa have their origin inferred in Africa, with postulated dispersal events to the Mediterranean, the Canary Islands, Madagascar and Southeast Asia. The lineages within Spathelioideae evolved at a relatively constant diversification rate. However, abrupt changes in diversification rates are inferred from the beginning of the Miocene and during the Pliocene/Pleistocene. Main conclusions The geographical origin of Spathelioideae probably lies in Africa. The existence of a Neotropical lineage may be the result of a dispersal event at a time in the Late Cretaceous when South America and Africa were still quite close to each other (assuming that our age estimates are close to the actual ages), or by Gondwanan vicariance (assuming that our age estimates provide minimal ages only). Separation of land masses caused by sea level changes during the Pliocene and Pleistocene may have been triggers for speciation in the Caribbean genus Spathelia.     

West Wind Drift revisited: testing for directional dispersal in the Southern Hemisphere using event-based tree fitting

Aim Recent studies suggest that if constrained by prevailing wind or ocean currents dispersal may produce predictable, repeated distribution patterns. Dispersal mediated by the West Wind Drift (WWD) and Antarctic Circumpolar Current (AAC) has often been invoked to explain the floristic similarities of Australia, South America and New Zealand. If these systems have been important dispersal vectors then eastward dispersal – from Australia to New Zealand and the western Pacific to South America – is expected to predominate. We investigate whether phylogenies for Southern Hemisphere plant groups provide evidence of historical dispersal asymmetry and more specifically whether inferred asymmetries are consistent with the direction of the WWD/AAC. Location Southern Hemisphere. Methods We assembled a data set of 23 published phylogenies for plant groups that occur in New Zealand, Australia and/or South America. We used parsimony‐based tree fitting to infer the number and direction of dispersals within each group. Observed dispersal asymmetries were tested for significance against a distribution of expected values. Results Our analyses suggest that dispersal has played a major role in establishing present distributions and that there are significant patterns of asymmetry in Southern Hemisphere dispersal. Consistent with the eastward direction of the WWD/ACC, dispersal from Australia to New Zealand was inferred significantly more often than in the reverse direction. No significant patterns of dispersal asymmetry were found between the western Pacific landmasses and South America. However, eastward dispersal was more frequently inferred between Australia and South America, while for New Zealand–South American events westward dispersal was more common. Main conclusions Our results suggest that eastward circumpolar currents have constrained the dispersal of plants between Australia and New Zealand. However, the WWD/ACC appear to have had less of an influence on dispersal between the western Pacific landmasses and South America. This observation may suggest that differences in dispersal mechanism are important – direct wind or water dispersal vs. stepping‐stone dispersal along the Antarctic coast. While our analyses provide useful preliminary insights into dispersal asymmetry in the Southern Hemisphere we will need larger data sets and additional methodological advances in order to test fully these dispersal patterns and infer processes from phylogenetic data.     

Calibrated chronograms,fossils, outgroup relationships,and root priors: re‐examining the historical biogeography of Geraniales

We re‐examined the recent study by Palazzesi et al., (2012) published in the Biological Journal of the Linnean Society (107: 67–85), that presented the historical diversification of Geraniales using BEAST analysis of the plastid spacer trnL–F and of the non‐coding nuclear ribosomal internal transcribed spacers (ITS). Their study presented a set of new fossils within the order, generated a chronogram for Geraniales and other rosid orders using fossil‐based priors on five nodes, demonstrated an Eocene radiation of Geraniales (and other rosid orders), and argued for more recent (Pliocene–Pleistocene) and climate‐linked diversification of genera in the five recognized families relative to previous studies. As a result of very young ages for the crown of Geraniales and other rosid orders, unusual relationships of Geraniales to other rosids, and apparent nucleotide substitution saturation of the two gene regions, we conducted a broad series of BEAST analyses that incorporated additional rosid fossil priors, used more accepted rosid ordinal topologies, or altered the placement of one fossil Geraniales prior. Our results indicate that their ages are 20–50% too young owing to a combination of (1) strong nucleotide saturation of the DNA regions starting at 65 Mya, (2) lack of a root (rosid stem) or other rosid ordinal stem fossil‐based priors, (3) the inability of the two DNA regions (with alignment issues) to obtain a monophyletic Geraniales as well as reasonable relationships of Geraniales to other rosid orders, and (4) apparent issues with the nodal placement of a Pelargonium fossil. The Geraniales crown is much older (Campanian of the Cretaceous; 86 Mya), the posterior age distribution on all but two fossil nodes are well older than the priors, the placement of a Pelargonium‐like fossil is more likely at the crown rather than the stem, but their models of diversification within several clades linked to climatic and orogeny appear supported. We discuss a number of the inherent issues of relaxed‐clock dating and outline some ‘best practice’ approaches for such studies. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 29–49.     

Evolution and phylogeny of the New Zealand cicada genus Kikihia Dugdale (Homoptera: Auchenorrhyncha: Cicadidae) with special reference to the origin of the Kermadec and Norfolk Islands' species

Aim Determine the phylogeny and dispersal patterns of the cicada genus Kikihia in New Zealand and the origin of the Norfolk, Kermadec, and Chatham Island cicadas. Location New Zealand, Norfolk Island, Kermadec Islands and Chatham Island. Methods DNA sequences from 16 species and four soon to be described species of cicadas from New Zealand and Norfolk Island (Australia) were examined. A total of 1401 base pairs were analysed from whole genome extraction of three mitochondrial genes (cytochrome oxidase subunit II, ATPase6 and ATPase8). These DNA sequences were aligned and analysed using standard likelihood approaches to phylogenetic analysis. Dates of divergences between clades were determined using a molecular clock based on Bayesian statistics. Results Most species in the genus Kikihia diverged between 3 and 5 million years ago (Ma) coincident with a period of rapid mountain building in New Zealand. Cicada species on the Kermadec and Norfolk Islands invaded recently from New Zealand and are closely related to the New Zealand North Island species Kikihia cutora. Main conclusions Speciation in the genus Kikihia was likely due in large part to the appearance of new habitats associated with the rise of the Southern Alps, starting c. 5 Ma. Dispersal of Kikihia species within mainland New Zealand probably occurred gradually rather than through long‐distance jumps. However, invasion of Norfolk, the Kermadecs and Chatham Islands had to have occurred through long‐distance dispersal.     

The fauna of athalassic saline waters in Australia and the Altiplano of South America: comparisons and historical perspectives

Similarities and differences between the fauna of inland saline waters in Australia and on the Altiplano are explored and explanations sought.Elements common to both continents include the calanoid copepod genus Boeckella (B. triarticulata in Australia, B. poopoensis and B. meteoris in South America) and the cladoceran genus Daphniopsis. Salinity data for Altiplano lakes are given for six species of Boeckella and for Daphniopsis.Ostracods have adapted to the open water of saline lakes in Australia but not in South America, a difference that may reflect past differences in the degree of predation by birds. In South America, diatoms are grazed by the flamingos Phoenicoparrus andinus and P. jamesi, while in Australia the main diatom grazer is probably the aquatic oniscoid isopod Haloniscus searlei. However, at least four species of flamingos were present in Australia during the late Cenozoic and one or more of these may well have grazed diatoms. The extinction of diatom-grazing or carnivorous flamingos, or both, in Australia may have been factors in the unique colonization of inland saline waters by H. searlei.     

Warm–cold colonization: response of oaks to uplift of the Himalaya–Hengduan Mountains

Clarifying the relationship between distribution patterns of organisms and geological events is critical to understanding the impact of environmental changes on organismal evolution. Quercus sect. Heterobalanus is now distributed across the Himalaya–Hengduan Mountains (HHM) and warm lowland in East China, yet how the distribution patterns of this group changed in response to the HHM uplift remains largely unknown. This study examines the effect of tectonic events in the HHM region on the oaks, providing a biological perspective on the geological history of this region. Fifty‐six populations of Quercus sect. Heterobalanus were genotyped using four chloroplast DNA regions and nine nuclear simple sequence repeat loci to assess population structure and diversity, supplemented by molecular dating and ancestral area reconstructions. The underlying demographic dynamics were compared using ecological niche models of the species distributions during the last glacial maximum and the present. These analyses illustrate that Quercus sect. Heterobalanus diversified as the HHM uplifted and climatic cooling during the mid‐Miocene, colonizing the cold habitats from warm broadleaf mixed forests. Lineages in cold highlands and warm lowlands have diverged as a consequence of local adaptation to diverging climates since the late Miocene. Our results suggest that continuous uplift of the HHM in the late Miocene to early Pliocene accompanied by simultaneous cooling triggered the differentiation of oaks. The biogeography of Quercus sect. Heterobalanus illuminates the geological events responsible for the modern‐day HHM.