The breeding season of three species of kiwi (Apteryx) in captivity as determined from egg-laying dates

Records from kiwi ( Apteryx spp.) breeding centres were used to obtain laying dates of 403 North Island brown kiwi ( Apteryx australis mantelli ) eggs, 24 great spotted kiwi ( Apteryx haastii ) eggs and 25 little spotted kiwi ( Apteryx owenii ) eggs. North Island brown kiwi outdoors had an annual cycle of egg-laying with 88.5% of eggs laid from June to January. Eggs were laid in every month of the year, with fewest eggs in April (2.1 % of all eggs). The greatest number of eggs was laid in July (15.6% of all eggs), with a second peak of laying in October (13.5% of all eggs). Both the great spotted kiwi and little spotted kiwi had annual cycles of egg-laying, with the main egg-laying seasons being August-January (87.5% of eggs) for the former and July-December (96.0% of eggs) for the latter. The highest peaks of laying were in October and August, respectively. Kiwi eggs were considered to belong to the same clutch if they were laid no more than 40 days apart. Clutch size in the North Island brown and little spotted kiwi was most commonly one or two eggs. In the great spotted kiwi only clutches of one egg were found. The mean clutch size of North Island brown kiwi outdoors (1.51±0.05) did not differ from that of free-living North Island brown kiwi (1.33±0.09). Similarly, there was no difference in the mean clutch size of captive and free-living little spotted kiwi (1.39±0.14 and 1.10±0.07). The mean interval between eggs for kiwi outdoors did not differ between North Island brown kiwi and little spotted kiwi (27.4±0.5 and 30.7±1.8 days). The results indicate that all three species of kiwi maintain annual cycles of egg-laying in captivity, with the main egg-laying season being longer than in free-living birds.     

Ancient DNA recovers the origins of Māori feather cloaks

Feather cloaks ("kakahu"), particularly those adorned with kiwi feathers, are treasured items or "taonga" to the Māori people of "Aotearoa"/New Zealand. They are considered iconic expression of Māori culture. Despite their status, much of our knowledge of the materials used to construct cloaks, the provenance of cloaks, and the origins of cloak making itself, has been lost. We used ancient DNA methods to recover mitochondrial DNA sequences from 849 feather samples taken from 109 cloaks. We show that almost all (>99%) of the cloaks were constructed using feathers from North Island brown kiwi. Molecular sexing of nuclear DNA recovered from 92 feather cloak samples also revealed that the sex ratio of birds deviated from a ratio of 1:1 observed in reference populations. Additionally, we constructed a database of 185 mitochondrial control region DNA sequences of kiwi feathers comprising samples collected from 26 North Island locations together with data available from the literature. Genetic subdivision (G(ST)), nucleotide subdivision (N(ST)) and Spatial Analysis of Molecular Variants (SAMOVA) analyses revealed high levels of genetic structuring in North Island brown kiwi. Together with sequence data from previously studied ancient and modern kiwi samples, we were able to determine the geographic provenance of 847 cloak feathers from 108 cloaks. A surprising proportion (15%) of cloaks were found to contain feathers from different geographic locations, providing evidence of kiwi trading among Māori tribes or organized hunting trips into other tribal areas. Our data also suggest that the east of the North Island of New Zealand was the most prolific of all kiwi cloak making areas, with over 50% of all cloaks analyzed originating from this region. Similar molecular approaches have the potential to discover a wealth of lost information from artifacts of endemic cultures worldwide.     

Molecular and other biological evidence supports the recognition of at least three species of brown kiwi

The presence of morphologically crypticlineages within the threatened brown kiwi ofNew Zealand has confounded their taxonomy. Recent genetic studies (Herbert and Daugherty1994; Baker et al. 1995) revealed that atleast two phylogenetic species exist within thebrown kiwi, and suggested that further researchshould resolve the taxonomic problems. In thispaper we extend genetic analyses to includesequences from 58 brown kiwi representing fivephylogenetic lineages for four mitochondrialloci (control region, cytochrome b,ATPase 6 and ATPase 8). Major lineages ofbrown kiwi are shown to be reciprocallymonophyletic, and align with other biologicaldifferences in the ecology, behavior,morphology and parasites of kiwi. BecausemtDNA sequences of major lineages of kiwi arenot evolving in a clocklike manner, we used anew penalized likelihood method withrate-smoothing to date the divergence of NorthIsland brown kiwi and the geographicallyisolated Okarito population (rowi) at about 6.2Mya. These lineages diverged about 8.2 Myafrom the brown kiwi in Fiordland and Haast inthe southern part of the South Island, and arethus older than the species of spotted kiwi(5.8 Mya). Given their distinctness, long-termgeographical isolation, lack of hybridizationin introduced populations, and accumulation ofnew biological characters within theselineages, we hypothesize that reproductiveincompatibilities have probably arisen as well. We therefore recommend that these divergentlineages be formally recognized as fullspecies; Apteryx mantelli should bere-instated for the North Island brown kiwi,A. australis should be restricted to thetokoeka, and a new species A. rowiishould be erected to describe the rowi atOkarito. Tokoeka should be split into at leastthree conservation management units (Haast,Fiordland and Stewart Island [Rakiura]), butfurther research is required to determine theexact relationships and status of theselineages. Further investigation is alsorequired into the genetic structuring of theNorth Island brown kiwi to confirm conservationmanagement units on the North Island.     

Incubation Temperatures of Great Spotted Kiwi,

Incubation temperatures of the great spotted kiwi were studied by telemetry methods at the Otorohanga Zoological Society in October 1989. The male maintained the core temperature of the egg at about 28-31.8-degrees-C. When he emerged to feed at night, the female started to incubate. She did not have a brood patch, but could heat the egg to 28-28.5-degrees-C, sufficient for embryo growth. Some of the reasons why female great spotted kiwi might help with incubation are discussed. In cold, mountain environments, the energetic costs of incubation could be so high that males alone cannot meet them. The hypothesis predicts that there are also places in the North Island where female brown kiwi (Apteryx australis mantelli) should share in incubation.     

Ancient DNA and conservation: lessons from the endangered kiwi of New Zealand

Conservation genetics typically seeks to map the distribution of contemporary genetic variation across space, and to use the resulting genetic parameters to infer any likely short-term evolutionary consequences for rare and endangered species. Recent developments in the study of ancient DNA now enable the extension of genetic variation studies backwards in time and provide a context by which to interpret contemporary levels of genetic variation, in addition to any patterns of genetic change over time. Ancient DNA research can also help to determine past levels of genetic diversity, identify species' boundaries and reveal former ranges among morphologically cryptic taxa. Ancient DNA sequence data for the New Zealand kiwi (Apteryx spp.) are presented and we show that most ancient populations of rowi and tokoeka exhibited private mitochondrial DNA haplotypes. Moreover, we illustrate that the extinction of these populations has led to the loss of considerable genetic variation. We also use ancient DNA methodology to determine past species distributions for brown kiwi and great spotted kiwi whose bones are morphologically indistinguishable.     

Incubation behaviour and egg physiology of kiwi (

Incubation behaviour varies among the different taxa of kiwi. For North Island brown kiwi (Apteryx mantelli) and little spotted kiwi (A. owenii), only the male incubates the eggs, except for in the first week. Meanwhile, for Okarito brown kiwi (A. mantelli) and the tokoeka (A. australis), incubation is shared by both sexes. In addition, amongst southern tokoeka, family group members can assist with incubation to the extent that breeding males may take no part in incubation at all. This study shows that kiwi turn their eggs regularly, an observation that contradicts earlier findings. There is a mean incubation temperature of 36.5°C at the top of the egg but the bottom may be 10°C lower in North Island brown kiwi and 5-8°C lower in Okarito brown kiwi eggs. Experiments inducing water loss from fertile and infertile eggs show that the presence of an embryo does not influence rate of water loss. North Island brown kiwi, Okarito brown kiwi and tokoeka regularly have more than one clutch in a season, and some North Island brown kiwi females lay up to seven eggs each year.     

Genetic consequences of a century of protection: serial founder events and survival of the little spotted kiwi (Apteryx owenii)

We present the outcome of a century of post-bottleneck isolation of a long-lived species, the little spotted kiwi (Apteryx owenii, LSK) and demonstrate that profound genetic consequences can result from protecting few individuals in isolation. LSK were saved from extinction by translocation of five birds from South Island, New Zealand to Kapiti Island 100 years ago. The Kapiti population now numbers some 1200 birds and provides founders for new populations. We used 15 microsatellite loci to compare genetic variation among Kapiti LSK and the populations of Red Mercury, Tiritiri Matangi and Long Islands that were founded with birds from Kapiti. Two LSK native to D''Urville Island were also placed on Long Island. We found extremely low genetic variation and signatures of acute and recent genetic bottleneck effects in all four populations, indicating that LSK have survived multiple genetic bottlenecks. The Long Island population appears to have arisen from a single mating pair from Kapiti, suggesting there is no genetic contribution from D''Urville birds among extant LSK. The N_e/N_C ratio of Kapiti Island LSK (0.03) is exceptionally low for terrestrial vertebrates and suggests that genetic diversity might still be eroding in this population, despite its large census size.     

Role of predation in the decline of kiwi,

Kiwi have declined markedly in abundance and range since human settlement of New Zealand. Three of the four species are still extant in mainland forests, despite decades of co-existence with various introduced mammals. Little spotted kiwi is now probably confined to offshore islands. The role of introduced mammals in these population declines was evaluated by measuring the survival rates of adults, eggs and chicks of brown kiwi (A. mantelli) and great spotted kiwi (A. haastii) in mainland forests. Mortality rates of adults ranged from 5%-16% and did not differ significantly between species or sexes. Overall, 14 out of 209 adult kiwi died during 159.6 radio-tracking years. Predators definitely caused five of these deaths. Sixty-nine (68%) of 102 eggs from 77 nesting attempts by 48 pairs failed to hatch. Predators probably caused about 10% of egg failures. Only three of 49 chicks probably survived to adulthood, indicating a juvenile mortality rate of about 94%. Predators killed at least 8% of chicks, 45% of juveniles, and possibly as many as 60% of all young kiwi. Ferrets and dogs were the main predators of adult kiwi, possums and mustelids were the main egg predators, while stoats and cats were largely responsible for the deaths of young kiwi. Population models show that northern brown kiwi are currently declining at 5.8% per annum. This decline could be halted by cutting the current predation rates on young kiwi by about 34% to 33%.     

Nuclear microsatellite DNA markers for New Zealand kiwi (Apteryx spp.)

Kiwi (Apterygidae) is an endemic New Zealand avian family comprising five species whose conservation is actively managed. We present five polymorphic microsatellite DNA loci isolated from North Island brown kiwi (Apteryx mantelli). In addition, we demonstrate cross‐amplification, and in some cases, polymorphism, of these microsatellite DNA loci in four other kiwi species. Therefore, these markers may be broadly applicable to conservation genetic studies within this family.     

Characterisation of class II B MHC genes from a ratite bird,the little spotted kiwi (<Emphasis Type="Italic">Apteryx owenii</Emphasis>)

Major histocompatibility complex (MHC) genes are important for vertebrate immune response and typically display high levels of diversity due to balancing selection from exposure to diverse pathogens. An understanding of the structure of the MHC region and diversity among functional MHC genes is critical to understanding the evolution of the MHC and species resilience to disease exposure. In this study, we characterise the structure and diversity of class II MHC genes in little spotted kiwi Apteryx owenii, a ratite bird representing the basal avian lineage (paleognaths). Results indicate that little spotted kiwi have a more complex MHC structure than that of other non-passerine birds, with at least five class II MHC genes, three of which are expressed and likely to be functional. Levels of MHC variation among little spotted kiwi are extremely low, with 13 birds assayed having nearly identical MHC genotypes (only two genotypes containing four alleles, three of which are fixed). These results suggest that recent genetic drift due to a species-wide bottleneck of at most seven birds has overwhelmed past selection for high MHC diversity in little spotted kiwi, potentially leaving the species highly susceptible to disease.     

Microevolution of the mitochondrial DNA control region in the Japanese brown bear (Ursus arctos) population.

We investigated nucleotide sequences of the mitochondrial DNA control region to describe natural genetic variations and to assess the relationships between subpopulations of the brown bear Ursus arctos on Hokkaido Island, Japan. Using the polymerase chain reaction product-direct sequencing technique, partial sequences (about 930 bases) of the control region were determined for 56 brown bears sampled throughout Hokkaido Island. A sequence alignment revealed that the brown bear control region included a variable sequence on the 5' side and a repetitive region on the 3' side. Phylogenetic trees reconstructed from the 5' variable region (696-702 bases) exhibited 17 haplotypes, which were clustered into three groups (Clusters A, B, and C). The distribution of each group did not overlap with those of the others, and the three different areas were located in separate mountainous forests of Hokkaido Island. Furthermore, most of the phylogenetically close haplotypes within each group were distributed geographically close to each other. In addition, the 3' repetitive region (arrays of 10 bases) exhibited a much faster mutation rate than the 5' variable region, resulting in heteroplasmy. Such mitochondrial DNA divergence in each group could have occurred after the brown bears migrated from the continent to Hokkaido and became fixed in the different areas.     

Population genetic subdivision in the New Zealand greenshell mussel (Perna canaliculus) inferred from single-strand conformation polymorphism analysis of mitochondrial DNA

Single-strand conformation polymorphism (SSCP) analysis of the NADH IV region of the mitochondrial DNA (mtDNA) molecule in greenshell mussels (Perna canaliculus) indicated strong population genetic structuring in this endemic New Zealand species. A northern and a southern group were differentiated by frequency shifts in common haplotypes and by the occurrence of a unique southern haplotype at approximately 20% frequency. This split occurred south of Cook Strait (the body of water between the North and the South Island) at approximately 42 degrees S latitude. Northern populations were less genetically diverse than southern populations and mussels from the west coast of the South Island were most distinct from northern mussels. We hypothesize that the unique haplotype VIII originated in the lower South Island, and that its spread northwards was obstructed by the opening of Cook Strait approximately 15 000-16 000 years ago and the subsequent establishment of present-day surface water circulation patterns in Greater Cook Strait. We suggest that present-day strong tidal flows and turbulent mixing of water masses in Cook Strait, and intense up-welling on the east and west coasts in this region, represent a barrier to gene flow between mussels located in the North Island and northern South Island vs. mussels in most of the South Island and Stewart Island.     

Molecular characterization of Babesia kiwiensis from the brown kiwi (Apteryx mantelli)

To further investigate the recently described avian piroplasm, Babesia kiwiensis, blood samples were collected from 13 wild-caught and 8 zoo-captive brown kiwi (Apteryx mantelli) and screened for the presence of piroplasm DNA using a nested-polymerase chain reaction (PCR) targeting the 18S rRNA gene of most members of Piroplasmida. All captive birds gave a negative PCR result, while 12 wild-caught birds were PCR positive. The nearly full-length 18S rRNA gene for B. kiwiensis was sequenced. Upon phylogenetic analysis, it was found to belong to the babesid group of piroplasms and was ancestral, yet genetically similar, to the Babesia canis-related species. An insight into the current taxonomy of the avian piroplasms is also given. An Ixodes anatis tick collected from 1 of the North Island brown kiwi was also screened using PCR and was found to be positive for B. kiwiensis DNA.     

Multilocus assignment analyses reveal multiple units and rare migration events in the recently expanded yellow-eyed penguin (Megadyptes antipodes)

The identification of demographically independent populations and the recognition of management units have been greatly facilitated by the continuing advances in genetic tools. Managements units now play a key role in short-term conservation management programmes of declining species, but their importance in expanding populations receives comparatively little attention. The endangered yellow-eyed penguin ( Megadyptes antipodes ) expanded its range from the subantarctic to New Zealand's South Island a few hundred years ago and this new population now represents almost half of the species' total census size. This dramatic expansion attests to M. antipodes' high dispersal abilities and suggests the species is likely to constitute a single demographic population. Here we test this hypothesis of panmixia by investigating genetic differentiation and levels of gene flow among penguin breeding areas using 12 autosomal microsatellite loci along with mitochondrial control region sequence analyses for 350 individuals. Contrary to our hypothesis, however, the analyses reveal two genetically and geographically distinct assemblages: South Island vs. subantarctic populations. Using assignment tests, we recognize just two first-generation migrants between these populations (corresponding to a migration rate of < 2%), indicating that ongoing levels of long-distance migration are low. Furthermore, the South Island population has low genetic variability compared to the subantarctic population. These results suggest that the South Island population was founded by only a small number of individuals, and that subsequent levels of gene flow have remained low. The demographic independence of the two populations warrants their designation as distinct management units and conservation efforts should be adjusted accordingly to protect both populations.     

Extensive intraspecific genetic diversity of a freshwater crayfish in a biodiversity hotspot

1. The freshwater crayfish Cherax dispar (Decapoda: Parastacidae) inhabits coastal regions and islands of South East Queensland, Australia. We hypothesised that populations of C. dispar on different islands would be more genetically divergent from each other than populations from different drainages within the same island or on the mainland. 2. Phylogenetic and phylogeographic analyses were conducted on two mitochondrial genes (cytochrome oxidase subunit I & 16S ribosomal DNA) and one nuclear gene (Internal Transcribed Spacer region 2). Phylogeographic patterns were compared with those for other freshwater organisms in the area. 3. Deep genetic divergences were found within C. dispar, including four highly divergent (up to 20%) clades. The geographic distribution of each of the clades revealed strong latitudinal structuring along the coast rather than structuring among the islands. The high genetic divergence observed among the C. dispar clades was estimated to have pre‐dated island formation and may represent ancient river drainage patterns. 4. A restricted distribution was observed for the most divergent clade, which was discovered only on two of the sand islands (North Stradbroke Island and Moreton Island). Furthermore, strong phylogeographic structuring was observed within this clade on North Stradbroke Island, where no haplotypes were shared between samples from opposite sides of the island. This low connectivity within the island supports the idea that C. dispar rarely disperse terrestrially (i.e. across watersheds).     

Genetic structure and phylogeography of European catfish (Silurus glanis) populations

The genetic structure of Silurus glanis (Europe's largest freshwater fish species) across most of its natural distribution was investigated using 10 microsatellite loci. The revealed levels of genetic diversity were much higher than previous allozyme and restriction fragment length polymorphism mitochondrial DNA analyses had shown; relative levels of variability among populations were however, in good agreement with the previous studies. Populations from large basins (Volga and Danube rivers) were the most polymorphic, while samples from the smaller Greek rivers, which are more prone to genetic bottleneck, exhibited the lowest levels of genetic diversity. Microsatellite multilocus genotyping permitted the assignment of individual fish to their population of origin with a score as high as 98.3%. Despite the great genetic differentiation of S. glanis populations, no consistent pattern of geographical structuring was revealed, in contrast to previous studies of European freshwater fish species. A model of isolation by distance seems more probable and a hypothesis of recent dispersion from only one glacial refugium is proposed. The discovery of the highest levels of microsatellite and mitochondrial diversity in the Volga sample and the presence of river connections, during the Pleistocene, between this area and all major areas of the present catfish distribution, place this refugium around the Ponto-Caspian region. Combining these data with those from previous studies, a number of markers are now available to monitor wild and hatchery populations even at the individual level.     

Closing the gap: Avian lineage splits at a young,narrow seaway imply a protracted history of mixed population response

The evolutionary significance of spatial habitat gaps has been well recognized since Alfred Russel Wallace compared the faunas of Bali and Lombok. Gaps between islands influence population structuring of some species, and flightless birds are expected to show strong partitioning even where habitat gaps are narrow. We examined the population structure of the most numerous living flightless land bird in New Zealand, Weka (Gallirallus australis). We surveyed Weka and their feather lice in native and introduced populations using genetic data gathered from DNA sequences of mitochondrial genes and nuclear β‐fibrinogen and five microsatellite loci. We found low genetic diversity among extant Weka population samples. Two genetic clusters were evident in the mtDNA from Weka and their lice, but partitioning at nuclear loci was less abrupt. Many formerly recognized subspecies/species were not supported; instead, we infer one subspecies for each of the two main New Zealand islands. Although currently range restricted, North Island Weka have higher mtDNA diversity than the more wide‐ranging southern Weka. Mismatch and neutrality statistics indicate North Island Weka experienced rapid and recent population reduction, while South Island Weka display the signature of recent expansion. Similar haplotype data from a widespread flying relative of Weka and other New Zealand birds revealed instances of North Island—South Island partitioning associated with a narrow habitat gap (Cook Strait). However, contrasting patterns indicate priority effects and other ecological factors have a strong influence on spatial exchange at this scale.     

Genetic structure and diversity of a rare woodland bat, <Emphasis Type="Italic">Myotis bechsteinii</Emphasis>: comparison of continental Europe and Britain

The Bechstein’s bat (Myotis bechsteinii) is a rare sedentary bat considered to be highly reliant on the presence of ancient woodland. Understanding the genetic connectivity and population structure of such elusive mammals is important for assessing their conservation status. In this study, we report the genetic diversity and structure of M. bechsteinii across Britain and Europe. Assessments were made using 14 microsatellite markers and a 747 bp region of the mitochondrial cytochrome b gene. Nuclear DNA (microsatellites) showed high levels of genetic diversity and little inbreeding across the species range, though genetic diversity was slightly lower in Britain than in mainland Europe. Bayesian and spatial PCA analysis showed a clear separation between the British and European sites. Within Europe, the Italian population south of the Alps was isolated from the other sites. In Britain, there was genetic structuring between the northern and southern part of the geographical range. Despite there being little genetic divergence in mitochondrial DNA (mtDNA) sequences throughout most of Europe, the mtDNA patterns in Britain confirmed this separation of northern and southern populations. Such genetic structuring within Britain—in the absence of any obvious physical barriers—suggests that other factors such as land-use may limit gene-flow.     

Glacial refugia in a maritime temperate climate: Cicada (Kikihia subalpina) mtDNA phylogeography in New Zealand

Understanding the biological significance of Pleistocene glaciations requires knowledge of the nature and extent of habitat refugia during glacial maxima. An opportunity to examine evidence of glacial forest refugia in a maritime, Southern Hemisphere setting is found in New Zealand, where the extent of Pleistocene forests remains controversial. We used the mitochondrial phylogeography of a forest-edge cicada ( Kikihia subalpina ) to test the hypothesis that populations of this species survived throughout South Island during the Last Glacial Maximum. We also compared mitochondrial DNA phylogeographic patterns with male song patterns that suggest allopatric divergence across Cook Strait. Cytochrome oxidase I and II sequences were analyzed using network analysis, maximum-likelihood phylogenetic estimation, Bayesian dating and Bayesian skyline plots. K. subalpina haplotypes from North Island and South Island form monophyletic clades that are concordant with song patterns. Song divergence corresponds to approximately 2% genetic divergence, and Bayesian dating suggests that the North Island and South Island population-lineages became isolated around 761 000 years bp . Almost all South Island genetic variation is found in the north of the island, consistent with refugia in Marlborough Sounds, central Nelson and northwest Nelson. All central and southern South Island and Stewart Island haplotypes are extremely similar to northern South Island haplotypes, a 'northern richness/southern purity' pattern that mirrors genetic patterns observed in many Northern Hemisphere taxa. Proposed southern South Island forest habitat fragments may have been too small to sustain populations of K. subalpina , and/or they may have harboured ecological communities with no modern-day analogues.     

Phylogeography of two New Zealand lizards: McCann's skink (Oligosoma maccanni) and the brown skink (O. zelandicum)

The New Zealand skink fauna has proven to be an ideal taxonomic group in which to examine the impact of climatic and geological processes on the evolution of the New Zealand biota since the Pliocene. Here we examine the phylogeography of McCann's skink (Oligosoma maccanni) in order to gain insight into the relative contribution of Pliocene and Pleistocene processes on patterns of genetic structure in the South Island biota, and investigate the phylogeography of the brown skink (O. zelandicum) to examine whether Cook Strait landbridges facilitated geneflow between the North and South Islands in the late-Pleistocene. We obtained mitochondrial DNA sequence data (ND2 and ND4; 1282bp) from across the range of both species. We examined the phylogeographic patterns evident in each species using Neighbour-Joining, Maximum Likelihood and Bayesian methods. We found substantial phylogeographic structure within O. maccanni, with seven distinct clades identified. Divergences among clades are estimated to have occurred during the Pliocene. Populations in the Otago/Southland region (south of the Waitaki River valley) formed a well-supported lineage within O. maccanni. A substantial genetic break was evident between populations in east and west Otago, either side of the Nevis-Cardrona fault system, while north-south genetic breaks were evident within the Canterbury region. Within-clade divergences in O. maccanni appear to have occurred during the mid- to late-Pleistocene. Shimodaira-Hasegawa topology tests indicated that the 'Garston' skink is not genetically distinct from O. maccanni. There was only relatively minor phylogeographic structure within O. zelandicum, with divergences among populations occurring during the mid- to late-Pleistocene. Our genetic data supports a single colonisation of the North Island by O. zelandicum from the South Island, with the estimated timing of this event (0.46mya) consistent with the initial formation of Cook Strait.