Soft-tissue specimens from pre-European extinct birds of New Zealand

We provide the first complete review of soft tissue remains from New Zealand birds that became extinct prior to European settlement (c. AD 1800). These rare specimens allow insights into the anatomy and appearance of the birds that are not attainable from bones. Our review includes previously unpublished records of ‘lost’ specimens, and descriptions of recently discovered specimens such as the first evidence of soft tissues from the South Island goose (Cnemiornis calcitrans). Overall, the soft tissue remains are dominated by moa (with specimens from each of the six genera), but also include specimens from Finsch's duck (Chenonetta finschi) and the New Zealand owlet-nightjar (Aegotheles novaezealandiae). All desiccated soft tissue specimens that have radiocarbon or stratigraphic dates are late Holocene in age, and most have been found in the semi-arid region of Central Otago

High-resolution coproecology: Using coprolites to reconstruct the habits and habitats of New Zealand’s extinct upland Moa (Megalapteryx didinus)

Knowledge about the diet and ecology of extinct herbivores has important implications for understanding the evolution of plant defence structures, establishing the influences of herbivory on past plant community structure and composition, and identifying pollination and seed dispersal syndromes. The flightless ratite moa (Aves: Dinornithiformes) were New Zealand's largest herbivores prior to their extinction soon after initial human settlement. Here we contribute to the knowledge of moa diet and ecology by reporting the results of a multidisciplinary study of 35 coprolites from a subalpine cave (Euphrates Cave) on the South Island of New Zealand. Ancient DNA analysis and radiocarbon dating revealed the coprolites were deposited by the extinct upland moa (Megalapteryx didinus), and span from at least 6,368±31 until 694±30 ¹⁴C years BP; the approximate time of their extinction. Using pollen, plant macrofossil, and ancient DNA analyses, we identified at least 67 plant taxa from the coprolites, including the first evidence that moa fed on the nectar-rich flowers of New Zealand flax (Phormium) and tree fuchsia (Fuchsia excorticata). The plant assemblage from the coprolites reflects a highly-generalist feeding ecology for upland moa, including browsing and grazing across the full range of locally available habitats (spanning southern beech (Nothofagus) forest to tussock (Chionochloa) grassland). Intact seeds in the coprolites indicate that upland moa may have been important dispersal agents for several plant taxa. Plant taxa with putative anti-browse adaptations were also identified in the coprolites. Clusters of coprolites (based on pollen assemblages, moa haplotypes, and radiocarbon dates), probably reflect specimens deposited at the same time by individual birds, and reveal the necessity of suitably large sample sizes in coprolite studies to overcome potential biases in diet interpretation

Evaluating the character and preservation of DNA within allophane clusters in buried soils on Holocene tephras, northern New Zealand

Clay minerals possess sorptive capacities for organic and inorganic matter, including DNA (Lorenz and Wackernagel, 1994), and hence reduce the utilization and degradation of organic matter or DNA by microorganisms. Buried allophane-rich soils on tephras (volcanic-ash beds) on the North Island, dated using tephrochronology, provide a valuable paleobiological ‘laboratory’ for studying the preservation of ancient DNA (aDNA) (Haile et al., 2007). Allophane comprises Al-rich nanocrystalline spherules ~3.5-5 nm in diameter (Fig. 1) with extremely large surface areas (up to 1000 m2 g-1). Moreover, allophanic soils are strongly associated with organic matter (Parfitt, 2009), and so we hypothesize that allophane also plays an important role for DNA protection within such soils

High-Resolution Coproecology: Using Coprolites to Reconstruct the Habits and Habitats of New Zealand’s Extinct Upland Moa (Megalapteryx didinus)

Knowledge about the diet and ecology of extinct herbivores has important implications for understanding the evolution of plant defence structures, establishing the influences of herbivory on past plant community structure and composition, and identifying pollination and seed dispersal syndromes. The flightless ratite moa (Aves: Dinornithiformes) were New Zealand’s largest herbivores prior to their extinction soon after initial human settlement. Here we contribute to the knowledge of moa diet and ecology by reporting the results of a multidisciplinary study of 35 coprolites from a subalpine cave (Euphrates Cave) on the South Island of New Zealand. Ancient DNA analysis and radiocarbon dating revealed the coprolites were deposited by the extinct upland moa (Megalapteryx didinus), and span from at least 6,368±31 until 694±30 14C years BP; the approximate time of their extinction. Using pollen, plant macrofossil, and ancient DNA analyses, we identified at least 67 plant taxa from the coprolites, including the first evidence that moa fed on the nectar-rich flowers of New Zealand flax (Phormium) and tree fuchsia (Fuchsia excorticata). The plant assemblage from the coprolites reflects a highly-generalist feeding ecology for upland moa, including browsing and grazing across the full range of locally available habitats (spanning southern beech (Nothofagus) forest to tussock (Chionochloa) grassland). Intact seeds in the coprolites indicate that upland moa may have been important dispersal agents for several plant taxa. Plant taxa with putative anti-browse adaptations were also identified in the coprolites. Clusters of coprolites (based on pollen assemblages, moa haplotypes, and radiocarbon dates), probably reflect specimens deposited at the same time by individual birds, and reveal the necessity of suitably large sample sizes in coprolite studies to overcome potential biases in diet interpretation

History Repeats: Large Scale Synchronous Biological Turnover in Avifauna From the Plio-Pleistocene and Late Holocene of New Zealand

New Zealand's unique biodiversity is the product of at least 55 million years of geographic isolation, supplemented by persistent transoceanic migration. Palaeontological and genetic evidence suggest most New Zealand avifauna has colonized from Australia. We synthesize evolutionary genetic studies to show a previously unrecognized clustering of divergence times in Australian and New Zealand bird species pairs, across the avian phylogeny at the beginning of the Pleistocene, around 2.5 million years ago. The timing coincides with major climatic and vegetation changes with the initiation of the Plio-Pleistocene glacial cycles. Recent anthropogenic impacts and environmental modifications are replicating in some important ways Pleistocene glacial landscapes, resulting in a new wave of avian “native invaders” into New Zealand

Characterizing porous microaggregates and soil organic matter sequestered in allophanic paleosols on Holocene tephras using synchrotron-based X-ray microscopy and spectroscopy

Allophanic tephra-derived soils can sequester sizable quantities of soil organic matter (SOM). However, no studies have visualized the fine internal porous structure of allophanic soil microaggregates, nor studied the carbon structure preserved in such soils or paleosols. We used synchrotron radiation-based transmission X-ray microscopy (TXM) to perform 3D-tomography of the internal porous structure of dominantly allophanic soil microaggregates, and carbon near-edge X-ray absorption fine-structure (C NEXAFS) spectroscopy to characterize SOM in ≤ 12,000-year-old tephra-derived allophane-rich (with minor ferrihydrite) paleosols. The TXM tomography showed a vast network of internal, tortuous nano-pores within an allophanic microaggregate comprising nanoaggregates. SOM in the allophanic paleosols at four sites was dominated by carboxylic/carbonyl functional groups with subordinate quinonic, aromatic, and aliphatic groups. All samples exhibited similar compositions despite differences between the sites. That the SOM does not comprise specific types of functional groups through time implies that the functional groups are relict. The SOM originated at the land/soil surface: ongoing tephra deposition (intermittently or abruptly) then caused the land-surface to rise so that the once-surface horizons were buried more deeply and hence became increasingly isolated from inputs by the surficial/modern organic cycle. The presence of quinonic carbon, from biological processes but vulnerable to oxygen and light, indicates the exceptional protection of SOM and bio-signals in allophanic paleosols, attributable both to the porous allophane (with ferrihydrite) aggregates that occlude the relict SOM from degradation, and to rapid burial by successive tephra-fallout, as well as strong Al-organic chemical bonding. TXM and C NEXAFS spectroscopy help to unravel the fine structure of soils and SOM and are of great potential for soil science studies

A new method to extract and purify DNA from allophanic soils and paleosols, and potential for paleoenvironmental reconstruction and other applications

Andisols, developed from late-Quaternary tephra (volcanic ash) deposits and dominated by the nanocrystalline aluminosilicate, allophane, contain large stores of organic matter and are potential reservoirs for DNA. However, DNA recovery from Andisols and other allophane-bearing soils has been difficult and inefficient because of strong chemical bonding between DNA and both allophane and organic matter, and also because much DNA can be encased and physically protected in nanopores in allophane nano/microaggregates. We have therefore developed a new two-step DNA isolation method for allophanic soils and buried paleosols, including those low in clay, which circumvents these problems. The method centres on (1) releasing mainly microbial DNA, and extracellular (unbound) DNA, using an alkaline phosphate buffer (“Rai’s lysis buffer”) that blocks re-adsorption sites on the allophanic materials, and (2) the novel application of acidified ammonium oxalate (Tamm’s reagent) to dissolve the allophane and to release DNA which had been chemically-bound and also which had been protected within nanopores. Ammonium oxalate has not previously been applied to soil DNA extraction. DNA yields up to 44.5 µg g-1 soil (oven-dry basis) were obtained from three field-moist natural allophanic soil samples from northern New Zealand using this two-step method. Following extraction, we evaluated different DNA purification methods. Gel electrophoresis of the extracted DNA followed by gel purification of the DNA from the agarose gel, despite some DNA loss, was the only purification method that removed sufficient humic material for successful DNA amplification using the polymerase chain reaction (PCR) of multiple gene regions. Sequencing of PCR products obtained from a buried allophanic paleosol at 2.2-m depth on a sandy Holocene tephra yielded endemic and exotic plants that differed from the European grasses growing currently on the soil’s surface. This difference suggests that the DNA extraction method is able to access (paleo)environmental DNA derived from previous vegetation cover. Our DNA extraction and purification method hence may be applied to Andisols and allophane-bearing paleosols, potentially offering a means to isolate paleoenvironmental DNA and thus facilitate reconstruction of past environments in volcanic landscapes, datable using tephrochronology, and also aid biodiversity understanding of andic soils and paleosols

Rapid radiation of Southern Ocean shags in response to receding sea ice

Understanding how natural populations respond to climatic shifts is a fundamental goal of biological research in a fast-changing world. The Southern Ocean represents a fascinating system for assessing large-scale climate-driven biological change, as it contains extremely isolated island groups within a predominantly westerly, circumpolar wind and current system. Blue-eyed shags represent a paradoxical seabird radiation—a circumpolar distribution implies strong dispersal capacity yet their species-rich nature suggests local adaptation and isolation. Here we attempt to resolve this paradox in light of the history of repeated cycles of climate change in the Southern Ocean

Ancient and contemporary DNA reveal a pre-human decline but no population bottleneck associated with recent human persecution in the kea (Nestor notabilis).

The impact of population bottlenecks is an important factor to consider when assessing species survival. Population declines can considerably limit the evolutionary potential of species and make them more susceptible to stochastic events. New Zealand has a well documented history of decline of endemic avifauna related to human colonization. Here, we investigate the genetic effects of a recent population decline in the endangered kea (Nestor notabilis). Kea have undergone a long-lasting persecution between the late 1800s to 1970s where an estimated 150,000 kea were culled under a governmental bounty scheme. Kea now number 1,000-5,000 individuals in the wild and it is likely that the recent population decline may have reduced the genetic diversity of the species. Comparison of contemporary (n = 410), historical (n = 15) and fossil samples (n = 4) showed a loss of mitochondrial diversity since the end of the last glaciation (Otiran Glacial) but no loss of overall genetic diversity associated with the cull. Microsatellite data indicated a recent bottleneck for only one population and a range-wide decline in Ne dating back some 300 - 6,000 years ago, a period predating European arrival in NZ. These results suggest that despite a recent human persecution, kea might have experienced a large population decline before stabilizing in numbers prior to human settlement of New Zealand in response to Holocene changes in habitat distribution. Our study therefore highlights the need to understand the respective effects of climate change and human activities on endangered species dynamics when proposing conservation guidelines

Did interaction between human pressure and Little Ice Age drive biological turnover in New Zealand?

Aim To test for simultaneous Holocene biogeographic turnover events in the New Zealand region. Specifically, we synthesize ancient DNA, radiocarbon data and archaeological data to assess the chronologies of late Holocene lineage extinction and replacement. Location Cool-temperate coastal ecosystems of New Zealand and the subantarctic. Methods We present new ancient DNA and radiocarbon data for New Zealand sea lions, and synthesize existing climatic, genetic and archaeological data, to test for synchronous megafaunal extinction and replacement events. The collated data include ancient DNA sequences from over 200 ancient sea lion and penguin specimens, in addition to 150 modern genetic samples. Results Our temporal genetic analyses show that, following human-driven extinction events, synchronous marine megafaunal replacement events occurred at around 1500 AD, coinciding with the Little Ice Age onset and an associated drastic human demographic decline in southern New Zealand. Conclusions A combination of climatic and human demographic shifts likely facilitated northward expansion of subantarctic sea lion and penguin lineages, replacing extirpated mainland New Zealand marine megafauna. Broadly, the interaction between human pressure and late Holocene climatic change may explain multiple biological turnover events in the Southern Hemisphere.Funding was provided by the Royal Society of New Zealand Marsden Fund (UOO1112), the University of Otago and the Allan Wilson Centre