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Long-term changes in the life history and behaviour of seabirds during the non-breeding season can reflect shifts in environmental conditions. However, long-term marine studies are scarce, particularly on southern hemisphere seabirds. Here, we used moult scores from 86 Brown Skuas (Stercorarius antarcticus lonnbergi), a large predatory seabird breeding on the Chatham Islands, Aotearoa/New Zealand to model both the timing and duration of primary feather moult. In addition, we analysed stable isotope values (δ13C and δ15N) from 62 modern (2014–16) and ten museum tail feathers. These data provide insights into the non-breeding behaviour of Brown Skua. Interestingly, our results show that the primary feather moult occurred prior to birds departing the colony, starting on average on 2 January ± 5 days (SE). The average start of primary feather moult occurred five days prior to the end of breeding (7 January ± 10 days (SD)) and 42 days before the birds departed the colony (13 February ± 11 days (SD)). The average duration of primary feather moult was 189 ± 14 days (SE). Importantly, low δ13C values in four females suggested that tail feather moult might also occur while skuas are at the colony. There was no difference in tail feather δ13C and δ15N values between any pairwise comparison of modern and museum years. However, values of δ15N from tail feathers sampled in 2014 were different from those sampled in 2015 and 2016. This large annual variation in δ15N values from tail feathers over such a short period makes long-term comparisons difficult to interpret, particularly between years with low sample sizes. While the stable isotope analyses of tail feathers are informative, we recommend future studies of skuas sample the primary coverts rather than tail feathers.

We report new avian remains from the early Eocene Nanjemoy Formation of the Fisher/Sullivan site in Virginia, USA. The fossil material includes the first records of the Messelasturidae, Psittacoped...

We describe a new species of Polynesian sandpiper from Henderson Island, Prosobonia sauli sp. nov., based on multiple Holocene fossil bones collected during the Sir Peter Scott Commemorative Expedition to the Pitcairn Islands (1991–92). Prosobonia sauli is the only species of Prosobonia to be described from bone accumulations and extends the record of known extinct Polynesian sandpipers to four. It is readily differentiated from the extant Tuamotu Sandpiper P. parvirostris in several features of the legs and bill, implying ecological adaptations to different environments. The geographically nearest Prosobonia populations to Henderson Island were found on Mangareva, where it is now extinct. A previous record of a species of Prosobonia from Tubuai, Austral Islands, is here shown to belong to the Sanderling Calidris alba. Our analyses of newly sequenced genetic data, which include the mitochondrial genomes of P. parvirostris and the extinct Tahiti Sandpiper P. leucoptera, confidently resolve the position of Prosobonia as sister-taxon to turnstones and calidrine sandpipers. We present a hypothesis for the timing of divergence between species of Prosobonia and other scolopacid lineages. Our results further provide a framework to interpret the evolution of sedentary lineages within the normally highly migratory Scolopacidae.

SummaryCook’s Petrel Pterodroma cookii is an endemic New Zealand seabird that has experienced a large range decline since the arrival of humans and now only breeds on two offshore islands (Te Hauturu-o-Toi/Little Barrier Island and Whenua Hou/Codfish Island) at the extreme ends of its former distribution. Morphological, behavioural, and mitochondrial cytochrome oxidase 1 (CO1) sequence data led a previous study to recognise the two extant populations as distinct conservation management units. Here, we further examine the genetic relationship between the extant populations using two nuclear introns (β-fibint7 and PAX). Using one mitochondrial locus (CO1), we also investigate the past distribution of a single nucleotide polymorphism (SNP) that differentiates the modern populations using bone and museum skins sourced from within its former range across New Zealand’s North and South Islands. We found significant population genetic structure between the two extant Cook’s Petrel populations for one of the two nuclear introns (β-fibint7). The mitochondrial DNA CO1 analysis indicated that the SNP variant found in the Codfish Island population was formerly widely distributed across both the North and South Islands, whereas the Little Barrier Island variant was detected only in North Island samples. We argue that these combined data support the recognition of the extant populations as different subspecies. Previous names for these taxa exist, thus Cook’s Petrel from Little Barrier Island becomes Pterodroma cookii cookii and Cook’s Petrel from Codfish Island becomes P. c. orientalis. Furthermore, we suggest that both genetic and non-genetic data should be taken into consideration when planning future mainland translocations. Namely, any translocations on the South Island should be sourced from Codfish Island and future translocations on the North Island should continue to be sourced from Little Barrier Island only.

Tobias, Joseph A., Sheard, Catherine, Pigot, Alex L., Devenish, Adam J. M., Yang, Jingyi, Sayol, Ferran, Neate‐Clegg, Montague H. C., Alioravainen, Nico, Weeks, Thomas L., Barber, Robert A., Walkden, Patrick A., MacGregor, Hannah E. A., Jones, Samuel E. I., Vincent, Claire, Phillips, Anna G., Marples, Nicola M., Montaño‐Centellas, Flavia A., Leandro‐Silva, Victor, Claramunt, Santiago, Darski, Bianca, Freeman, Benjamin G., Bregman, Tom P., Cooney, Christopher R., Hughes, Emma C., Capp, Elliot J. R., Varley, Zoë K., Friedman, Nicholas R., Korntheuer, Heiko, Corrales‐Vargas, Andrea, Trisos, Christopher H., Weeks, Brian C., Hanz, Dagmar M., Töpfer, Till, Bravo, Gustavo A., Remeš, Vladimír, Nowak, Larissa, Carneiro, Lincoln S., Moncada R., Amilkar J., Matysioková, Beata, Baldassarre, Daniel T., Martínez‐Salinas, Alejandra, Wolfe, Jared D., Chapman, Philip M., Daly, Benjamin G., Sorensen, Marjorie C., Neu, Alexander, Ford, Michael A., Mayhew, Rebekah J., Fabio Silveira, Luis, Kelly, David J., Annorbah, Nathaniel N. D., Pollock, Henry S., Grabowska‐Zhang, Ada M., McEntee, Jay P., Carlos T. Gonzalez, Juan, Meneses, Camila G., Muñoz, Marcia C., Powell, Luke L., Jamie, Gabriel A., Matthews, Thomas J., Johnson, Oscar, Brito, Guilherme R. R., Zyskowski, Kristof, Crates, Ross, Harvey, Michael G., Jurado Zevallos, Maura, Hosner, Peter A., Bradfer‐Lawrence, Tom, Maley, James M., Stiles, F. Gary, Lima, Hevana S., Provost, Kaiya L., Chibesa, Moses, Mashao, Mmatjie, Howard, Jeffrey T., Mlamba, Edson, Chua, Marcus A. H., Li, Bicheng, Gómez, M. Isabel, García, Natalia C., Päckert, Martin, Fuchs, Jérôme, Ali, Jarome R., Derryberry, Elizabeth P., Carlson, Monica L., Urriza, Rolly C., Brzeski, Kristin E., Prawiradilaga, Dewi M., Rayner, Matt J., Miller, Eliot T., Bowie, Rauri C. K., Lafontaine, René‐Marie, Scofield, R. Paul, Lou, Yingqiang, Somarathna, Lankani, Lepage, Denis, Illif, Marshall, Neuschulz, Eike Lena, Templin, Mathias, Dehling, D. Matthias, Cooper, Jacob C., Pauwels, Olivier S. G., Analuddin, Kangkuso, Fjeldså, Jon, Seddon, Nathalie, Sweet, Paul R., DeClerck, Fabrice A. J., Naka, Luciano N., Brawn, Jeffrey D., Aleixo, Alexandre, Böhning‐Gaese, Katrin, Rahbek, Carsten, Fritz, Susanne A., Thomas, Gavin H., Schleuning, Matthias (2022): AVONET: morphological, ecological and geographical data for all birds. Ecology Letters 25 (3): 581-597, DOI: 10.1111/ele.13898, URL: http://dx.doi.org/10.1111/ele.13898

Fossil penguins from late Neogene sediments are important for understanding the origins of modern penguins. Recent advances in penguin taxonomy and geometric morphometrics allow us to revisit previ...

Mayr, G., De Pietri, V.L., Love, L., Mannering, A. & Scofield, R.P., 9 August 2019. Leg bones of a new penguin species from the Waipara Greensand add to the diversity of very large-sized Spheniscif...

The early Miocene charadriiform bird Becassius charadriioides De Pietri and Mayr, 2012, from the Saint-Gerand-le-Puy area in France, was originally described as a member of uncertain affinities within the shorebird clade Scolopaci (jacanas, seedsnipe, painted-snipe, sandpipers, and allies). Following a re-assessment of the material attributed to this taxon and in the context of a larger comparative sample of extinct and extant charadriiform birds, we conclude that it is a member of the Glareolidae (pratincoles and coursers). We also demonstrate that certain elements, such as the coracoid, which were only tentatively referred to B. charadriioides, are very likely to belong to this taxon. We describe for the first time a tarsometatarsus that we tentatively attribute to this species. Based on the morphology of the humerus and other elements, it is not possible to associate Becassius charadriioides with any extant lineage within Glareolidae; it displays a combination of morphological features that can be presumed to be ancestral to Glareolidae based on outgroup comparisons and on the distinctiveness of B. charadriioides among other glareolids. The referral of Becassius charadriioides to Glareolidae bridges a gap in the evolutionary history of the clade, attesting to the presence of members of this clade in Europe during the earliest Miocene. Additionally, we provide a review of the fossil record of Glareolidae and re-assess some of the oldest fossils to have been attributed to this group.

Bishop, P.J., Scofield, R.P. & Hocknull, S.A. 6 May 2019. The architecture of cancellous bone in the hindlimb of moa (Aves: Dinornithiformes), with implications for stance and gait.. The architectu...

Prior to human arrival in the 13th century, two large birds of prey were the top predators in New Zealand. In the absence of non-volant mammals, the extinct Haast's eagle (Hieraaetus moorei), the largest eagle in the world, and the extinct Eyles' harrier (Circus teauteensis) the largest harrier in the world, had filled ecological niches that are on other landmasses occupied by animals such as large cats or canines. The evolutionary and biogeographic history of these island giants has long been a mystery. Here we reconstruct the origin and evolution of New Zealand’s giant raptors using complete mitochondrial genome data. We show that both Eyles’ harrier and Haast’s eagle diverged from much smaller, open land adapted Australasian relatives in the late Pliocene to early Pleistocene. These events coincided with the development of open habitat in the previously densely forested islands of New Zealand. Our study provides evidence of rapid evolution of island gigantism in New Zealand’s extinct birds of prey. Early Pleistocene climate and environmental changes were likely to have triggered the establishment of Australian raptors into New Zealand. Our results shed light on the evolution of two of the most impressive cases of island gigantism in the world.

Aim: Correlative species distribution models (SDMs) are typically trained using only the contemporary distribution of species; however, recent records might reflect an incomplete description of a species' niche, limiting the reliability of predictions. SDMs linking fossil records have the potential to improve conservation decisions under human-induced climate change. Here, we built SDMs using presence records from contemporary and Holocene records to enable estimations of climatically suitable area under current and future climate scenarios. Location: Aotearoa New Zealand. Taxon: Tuatara, Sphenodon punctatus. Methods: For an evolutionary relict found in Aotearoa New Zealand, the tuatara (Sphenodon punctatus), we built SDMs using presence records from contemporary and Holocene records to estimate climatically suitable area under current and future climate scenarios. We also use our detailed knowledge of the Holocene distribution and remnant populations to examine niche shifts following the arrival of humans and associated introduction of mammalian predators. To build SDMs, we use presence records from four sources: (a) remnant populations, (b) radiocarbon-dated fossil deposits from the Holocene, (c) other fossil deposits containing tuatara bones of Holocene age and iv) islands from which tuatara are known or highly likely to have become extinct. Results: We found shifts in the niche of tuatara due to niche unfilling. Incorporating locations of Holocene deposits and/or all past locations in SDMs led to larger areas of climatically suitable area being identified compared to SDMs derived from remnant populations only. Using all presence records, under climate change projections for 2090, climatically suitable area increased slightly. However, many areas retain potential as translocation sites (e.g. northern South Island), some areas become unsuitable (e.g. inland Canterbury) and/or involve extrapolation into novel climates (e.g. Northland). Main Conclusion: SDMs incorporating locations of Holocene deposits and/or all past locations identified areas of critical habitat for tuatara under current and future climate scenarios, that would not have been identified using contemporary occurrences only. Our results highlight the need to consider past locations when assessing habitat suitability for conservation translocations, both for tuatara and other relict species.

The holotype specimen of Alexandronectes zealandiensis is analyzed using digital reconstruction based on CT scans. Additional information regarding internal anatomy or obscured details are added. Additional features include: ectopterygoid rhombic in shape with posterior end pointed, a feature shared only with Aristonectes quiriquinensis; pterygoid shows a high dorsal crest located anteriorly and laterally to the level of the basipterygoid process; posterior margin of parabasisphenoid medially notched in ventral view and surrounding anteriorly and laterally a midline pit. Two canals for XII nerve are present in the right exoccipital-opisthothic and only one in the left one. Supraoccipital with two medially curved ridges on its posterior surface. Additionally, the presence of a stapes is described for the first time in an aristonectine elasmosaurid. The inner ear labyrinth is described and compared with that of other plesiosaurs, and the floccular recess (osseous correlate of the floccular lobe of the cerebellum) is described for first time among elasmosaurids. This feature is probably related to the presence of a long neck and with predatory behavior as the floccular lobe (housed in the floccular recess) stabilized the head via the cervical musculature, and stabilized the retinal image during rotational head movements.

Migratory marine species cross political borders and enter the high seas, where the lack of an effective global management framework for biodiversity leaves them vulnerable to threats. Here, we combine 10,108 tracks from 5775 individual birds at 87 sites with data on breeding population sizes to estimate the relative year-round importance of national jurisdictions and high seas areas for 39 species of albatrosses and large petrels. Populations from every country made extensive use of the high seas, indicating the stake each country has in the management of biodiversity in international waters. We quantified the links among national populations of these threatened seabirds and the regional fisheries management organizations (RFMOs) which regulate fishing in the high seas. This work makes explicit the relative responsibilities that each country and RFMO has for the management of shared biodiversity, providing invaluable information for the conservation and management of migratory species in the marine realm., This project has received funding from the European Union’s Horizon 2020 Research and Innovation Programme under the Marie Skłodowska-Curie grant agreement no. 766417. Research made possible with the support, permissions, and funding granted by the following organizations, institutions, and grant agreements: Marine Conservation Program of DPIPWE Tasmania; American Bird Conservancy; Bureau of Ocean Energy Management; communities of Isla Mocha and Islas Juan Fernández; Corporación Nacional Forestal and Servicio Agrícola y Ganadería (Chile); Environment Canada; the National Fish and Wildlife Foundation; the National Geographic Society; ProDelphinus; and the Wallis Foundation; NSF grants DEB 9304579, DEB 9629539, DEB9806606, DEB0235818, and DEB 0842199 to D.J.A.; the National Geographic Society; the U.S. Fish and Wildlife Service; U.S. Geological Survey Ecosystems Mission Area, Wake Forest University; Max-Planck Society and State of Baden-Wuerttemberg Innovation funding; Colorado State University International Programs; Swiss Friends of Galapagos; the International Center for Tropical Ecology at University of Missouri-St. Louis; the Instituto Antartico Chileno (INACH) and the Australian Antarctic Division (AAD); LIFE “Marine IBAs IN Spain” (LIFE04NAT/ES/000049, 2004-2009) and LIFE+ INDEMARES (2009-2014); Sea World Research and Rescue Foundation Inc.; Holsworth Wildlife Research Endowment; and Winifred Violet Scott Trust; FCT-Portugal through projects (UIDB/04292/2020 and UIDP/04292/2020 and UIDP/50017/2020 and UIDB/50017/2020, granted to MARE and CESAM, respectively); the Falklands Islands Government; Natural Environmental Research Council (NERC) core funding to British Antarctic Survey Ecosystems Programme and Official Development Assistance Atlantic Islands project (NE/ R000 107/1); the New Zealand Department of Conservation; Ministry for Primary Industries; Ngāti Rehua Ngāti Wai ki Aotea; Falklands Island Conservation; University of Barcelona (APIF/2015, to M.C.-F.); the French Polar Institute (program IPEV n°109 to H.W.); Réserve Naturelle Nationale des Terres Australes Françaises; and the Zones Atelier Antarctique (LTSER France, CNRS-INEE); European funds through the European Commission Training and Mobility of Researchers Programme (ERBFMBICT983030); Spanish funds through the Ministerio de Ciencia y Tecnología (REN2002-01164/GLO), Ministerio de Educación y Ciencia (CGL2006-01315/BOS), Ministerio de Ciencia e Innovación (CGL2009-11278/BOS), and Ministerio de Economía y Competitividad (CGL2013-42585-P); Catalan funds through the Generalitat de Catalunya (2001SGR00091); and additional funding from SEO/BirdLife (programa Migra & proyecto LIFE+ Indemares), Fundación Banco Bilbao Vizcaya Argentaria (BIOCON04/099) and Fundación Biodiversidad (18PCA4328, 2012-2013); NSERC Discovery Grant and Government of Canada’s Program for International Polar Year to W.A.M.; and an ACAP AC Grant in 2013-14, predoctoral contract BES-2017-079874 of the Spanish Ministerio de Industria, Economía y Competitividad (to L.N.-H.); Spanish Foundation for Biodiversity and Spanish Ministry of Science grant ref. CGL2013-42203-R; the Pew Environment Group via the Pew Fellowship Award in Marine Conservation (to M.L.C.); National Research Foundation; South Africa and Oceans and Coasts; Department of Environment, Agriculture and Fisheries; Malta Seabird Project (LIFE10NAT/MT/090) co-funded by the LIFE program of the European Commission and the Maltese Ministry for the Environment, Sustainable Development and Climate Change, in partnership with the Royal Society for the Protection of Bird and the Portuguese Society for the Study of Birds; predoctoral contract BES-2014-068025 of the Spanish Ministerio de Industria, Economía y Competitividad (to V.M.-P.); Scientific Expert PIM initiative (Petites Iles de Méditerranée); the PIM initiative (Petites Iles de Méditerranée); the Tunisian Coastal Protection and Planning Agency (APAL); Ministry of the Environment, Japan; Funding by Fundación Ecocentro, Argentina; Wildlife Conservation Society, USA; and Ministerio de Ciencia, Tecnología e Innovación, Argentina; Centro Nacional Patagónico (CONICET), postdoctoral contracts by Beatriu de Pinós (2010-BP_A-00173), Juan de la Cierva (JCI-2009-05426), PLEAMAR (2017/2349), and Ramón y Cajal (RYC-2017-22055) programme (to R.R.); Seventh Framework Programme (Research Executive Agency of the European Commission, 618841, FP7-PEOPLE-2013-CIG); Fondation Total pour la Biodiversité (project: Trophic ecology and impacts of bycatch on the avifauna communities of Zembra archipelago); Agence de Protection et d’Aménagement du Littoral (APAL-Tunisia); Killam Postdoctoral fellowship from Dalhousie University; South African National Antarctic Programme; ACAP; Papahānaumokuākea Marine National Monument; NOAA; Japan Society for the Promotion of Science Kakenhi grant 19651100 and 15H02857; National Parks and Conservation Service (Mauritius) (to M.L.C.); IPEV Prog 109; and NASA. The use of trade, product, or firm names in this publication is for descriptive purposes only and does not imply endorsement by the U.S. government. The scientific results and conclusions, as well as any views or opinions expressed herein, are those of the authors and do not necessarily reflect those of the NOAA or the Department of Commerce. This communication reflects only the authors’ view, and the Research Executive Agency of the European Union is not responsible for any use that may be made of the information it contains. Z.Z. acknowledges funding from a predoctoral grant (APIF/2012) from the University of Barcelona. J.Ad. acknowledges funding from the U.S. Geological Survey Ecosystems Mission Area, U.S. Bureau of Ocean Energy Management, Pacific OCS Region.

1. While floating near the sea surface plastic debris interacts with a number of external factors, including many different organisms. Seabirds have the most extensive documented history of interactions with plastics, through ingestion, entanglement, and nest construction. 2. In the present study, eight seabird species from the South Pacific Ocean were used as a proxy to determine potential patterns of removal of marine plastic debris, and three hypotheses were tested in relation to their feeding habits and nesting areas. 3. Plastics from abiotic compartments (Chilean continental coast, South Pacific Gyre, and Rapa Nui beaches) and biotic compartments (surface‐feeding seabirds, diving seabirds, and nesting areas) were compared, according to their type, colour, shape, and density. 4. Continental beaches had a relatively wide range of colours and shapes, with many non‐buoyant plastics. Samples from the South Pacific Gyre (SPG) and Rapa Nui (Easter Island) beaches comprised mainly hard, rounded, buoyant, and white/grey plastics. 5. These results indicate that the composition of floating plastics from terrestrial sources changes during transport with oceanic currents, reducing the proportion of prey‐like plastics present in the subtropical gyres. 6. The stomach contents of surface‐feeding and diving seabirds were dominated by hard, white/grey, and round plastic items, similar to plastics from the SPG, suggesting non‐selective (accidental or secondary) ingestion. 7. Nesting areas had a more variable composition of brightly coloured plastics, suggesting a pattern of selective removal of plastics by seabirds, probably from oceanic sources. 8. The present study reveals extensive interactions of seabirds with plastics on a broader scale, which is highly relevant given that the impacts of plastics on seabirds are increasing worldwide, compromising their efficient conservation.

New Zealand is a globally significant hotspot for seabird diversity, but the sparse fossil record for most seabird lineages has impeded our understanding of how and when this hotspot developed. Here, we describe multiple exceptionally well-preserved specimens of a new species of penguin from tightly dated (3.36–3.06 Ma) Pliocene deposits in New Zealand. Bayesian and parsimony analyses place Eudyptes atatu sp. nov. as the sister species to all extant and recently extinct members of the crested penguin genus Eudyptes . The new species has a markedly more slender upper beak and mandible compared with other Eudyptes penguins. Our combined evidence approach reveals that deep bills evolved in both crested and stiff-tailed penguins ( Pygoscelis ) during the Pliocene. That deep bills arose so late in the greater than 60 million year evolutionary history of penguins suggests that dietary shifts may have occurred as wind-driven Pliocene upwelling radically restructured southern ocean ecosystems. Ancestral area reconstructions using BioGeoBEARS identify New Zealand as the most likely ancestral area for total-group penguins, crown penguins and crested penguins. Our analyses provide a timeframe for recruitment of crown penguins into the New Zealand avifauna, indicating this process began in the late Neogene and was completed via multiple waves of colonizing lineages.

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1. The identification of geographic areas where the densities of animals are highest across their annual cycles is a crucial step in conservation planning. In marine environments, however, it can be particularly difficult to map the distribution of species, and the methods used are usually biased towards adults, neglecting the distribution of other life-history stages even though they can represent a substantial proportion of the total population. 2. Here we develop a methodological framework for estimating populationlevel density distributions of seabirds, integrating tracking data across the main life-history stages (adult breeders and non-breeders, juveniles and immatures). We incorporate demographic information (adult and juvenile/immature survival, breeding frequency and success, age at first breeding) and phenological data (average timing of breeding and migration) to weight distribution maps according to the proportion of the population represented by each life-history stage. 3. We demonstrate the utility of this framework by applying it to 22 species of albatrosses and petrels that are of conservation concern due to interactions with fisheries. Because juveniles, immatures and non-breeding adults account for 47%–81% of all individuals of the populations analysed, ignoring the distributions of birds in these stages leads to biased estimates of overlap with threats, and may misdirect management and conservation efforts. Population-level distribution maps using only adult distributions underestimated exposure to longline fishing effort by 18%–42%, compared with overlap scores based on data from all lifehistory stages. 4. Synthesis and applications. Our framework synthesizes and improves on previous approaches to estimate seabird densities at sea, is applicable for data-poor situations, and provides a standard and repeatable method that can be easily updated as new tracking and demographic data become available. We provide scripts in the R language and a Shiny app to facilitate future applications of our approach. We recommend that where sufficient tracking data are available, this framework be used to assess overlap of seabirds with at-sea threats such as overharvesting, fisheries bycatch, shipping, offshore industry and pollutants. Based on such an analysis, conservation interventions could be directed towards areas where they have the greatest impact on populations. Fundação para a Ciência e a Tecnologia - FCT; French Polar Institute IPEV; Falkland Islands Government info:eu-repo/semantics/publishedVersion

The arrival of humans in New Zealand around 750 years ago resulted in widespread faunal extinctions including the endemic Waitaha penguin (Megadyptes waitaha). Previously thought to have only bred on coastal South Island and Stewart Island, recent genetic reanalysis of prehistoric large penguin bones from the lower North Island indicates that the Waitaha penguin may have been a common resident. Here we synthesise previous studies and present new palaeontological and archaeological evidence to suggest that the Waitaha penguin was probably breeding in the lower North Island at the time of human arrival, and did not represent vagrant individuals from more southerly breeding colonies. The elimination of breeding Megadyptes from the North Island would add to the already significant avifaunal losses from New Zealand, of which the North Island suffered the greatest biodiversity loss after the arrival of humans.

Significance The mode and tempo of extinctions and extirpations after the first contact phase of human settlements is a widely debated topic. As the last major landmass to be settled by humans, New Zealand offers a unique lens through which to study interactions of people and biota. By analyzing ancient DNA from more than 5,000 nondiagnostic and fragmented bones from 38 subfossil assemblages, we describe species and patterns that have been missed by morphological approaches. We report the identification of five species of whale from an archaeological context in New Zealand and describe the prehistoric kākāpō population structure. Taken together, this study demonstrates insights into subsistence practices and extinction processes and demonstrates the value of genetic analyses of fossil assemblages., New Zealand’s geographic isolation, lack of native terrestrial mammals, and Gondwanan origins make it an ideal location to study evolutionary processes. However, since the archipelago was first settled by humans 750 y ago, its unique biodiversity has been under pressure, and today an estimated 49% of the terrestrial avifauna is extinct. Current efforts to conserve the remaining fauna rely on a better understanding of the composition of past ecosystems, as well as the causes and timing of past extinctions. The exact temporal and spatial dynamics of New Zealand’s extinct fauna, however, can be difficult to interpret, as only a small proportion of animals are preserved as morphologically identifiable fossils. Here, we conduct a large-scale genetic survey of subfossil bone assemblages to elucidate the impact of humans on the environment in New Zealand. By genetically identifying more than 5,000 nondiagnostic bone fragments from archaeological and paleontological sites, we reconstruct a rich faunal record of 110 species of birds, fish, reptiles, amphibians, and marine mammals. We report evidence of five whale species rarely reported from New Zealand archaeological middens and characterize extinct lineages of leiopelmatid frog (Leiopelma sp.) and kākāpō (Strigops habroptilus) haplotypes lost from the gene pool. Taken together, this molecular audit of New Zealand’s subfossil record not only contributes to our understanding of past biodiversity and precontact Māori subsistence practices but also provides a more nuanced snapshot of anthropogenic impacts on native fauna after first human arrival.

While known for over a decade to exist, fossil rails of the early Miocene (19–16 Ma) St Bathans Fauna, from the South Island of New Zealand, have not previously been described taxonomically or studied in detail. Here we use qualitative osteological features and analyse measurements from wing and leg bones to determine the number of taxa represented, their flight ability, and the presence and nature of sexual dimorphism within the identified taxa. We describe two new rail species in monospecific genera from the St Bathans Fauna: Priscaweka parvales gen. et sp. nov., which is extremely common, and Litorallus livezeyi gen. et sp. nov., a distinctly larger, uncommon species. Priscaweka parvales exhibited a significant degree of sexual dimorphism and was tiny, being the size of the extinct Chatham Island Rail Cabalus modestus. Both newly described species exhibit skeletal proportions and osteological features that indicate they had reduced wings and were flightless. These observations reveal that flightless ra...

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/., A new genus and species of fossil bat is described from New Zealand’s only pre-Pleistocene Cenozoic terrestrial fauna, the early Miocene St Bathans Fauna of Central Otago, South Island. Bayesian total evidence phylogenetic analysis places this new Southern Hemisphere taxon among the burrowing bats (mystacinids) of New Zealand and Australia, although its lower dentition also resembles Africa’s endemic sucker-footed bats (myzopodids). As the first new bat genus to be added to New Zealand’s fauna in more than 150 years, it provides new insight into the original diversity of chiropterans in Australasia. It also underscores the significant decline in morphological diversity that has taken place in the highly distinctive, semi-terrestrial bat family Mystacinidae since the Miocene. This bat was relatively large, with an estimated body mass of ~40 g, and its dentition suggests it had an omnivorous diet. Its striking dental autapomorphies, including development of a large hypocone, signal a shift of diet compared with other mystacinids, and may provide evidence of an adaptive radiation in feeding strategy in this group of noctilionoid bats.

New Zealand’s endemic King Shag ( Leucocarbo carunculatus ) has occupied only a narrow portion of the northeastern South Island for at least the past 240 years. However, pre-human Holocene fossil and archaeological remains have suggested a far more widespread distribution of the three Leucocarbo species (King, Otago, Foveaux) on mainland New Zealand at the time of Polynesian settlement in the late 13th Century CE. We use modern and ancient DNA, and morphometric and osteological analyses, of modern King Shags and Holocene fossil Leucocarbo remains to assess the pre-human distribution and taxonomic status of the King Shag on mainland New Zealand, and the resultant conservation implications. Our analyses show that the King Shag was formerly widespread around southern coasts of the North Island and the northern parts of the South Island but experienced population and lineage extinctions, and range contraction, probably after Polynesian arrival. This history parallels range contractions of other New Zealand seabirds. Conservation management of the King Shag should take into account this species narrow distribution and probable reduced genetic diversity. Moreover, combined genetic, morphometric and osteological analyses of prehistoric material from mainland New Zealand suggest that the now extinct northern New Zealand Leucocarbo populations comprised a unique lineage. Although these distinctive populations were previously assigned to the King Shag (based on morphological similarities and geographic proximity to modern Leucocarbo populations), we herein describe them as a new species, the Kohatu Shag ( Leucocarbo septentrionalis ). The extinction of this species further highlights the dramatic impacts Polynesians and introduced predators had on New Zealand’s coastal and marine biodiversity. The prehistoric presence of at least four species of Leucocarbo shag on mainland NZ further highlights its status as a biodiversity hotspot for Phalacrocoracidae.

This study is the first to utilize 30-cm resolution imagery from the WorldView-3 (WV-3) satellite to count wildlife directly. We test the accuracy of the satellite method for directly counting individuals at a well-studied colony of Wandering Albatross Diomedea exulans at South Georgia, and then apply it to the closely related Northern Royal Albatross Diomedea sanfordi, which is near-endemic to the Chatham Islands and of unknown recent population status due to the remoteness and limited accessibility of the colonies. At South Georgia, satellite-based counts were comparable to ground-based counts of Wandering Albatross nests, with a slight over-estimation due to the presence of non-breeding birds. In the Chatham Islands, satellite-based counts of Northern Royal Albatross in the 2015/2016 season were similar to ground-based counts undertaken on the Forty-Fours islands in 2009/2010, but much lower than ground-based counts undertaken on The Sisters islands in 2009/2010, which is of major conservation concern for this endangered albatross species. We conclude that the ground-breaking resolution of the newly available WV-3 satellite will provide a step change in our ability to count albatrosses and other large birds directly from space without disturbance, at potentially lower cost and with minimal logistical effort.

Recent genetic analyses have revealed evidence for the existence of two distinct lineages of Little Penguin (genus Eudyptula), one endemic to New Zealand (E. minor), and the other widespread along the coasts of Australia and south-east New Zealand (E. novaehollandiae). Here we present the first comprehensive morphometric analysis of complete Eudyptula skeletons. We show that variability in Eudyptula body and bill size is associated with sea surface temperature gradients, suggesting that thermoregulatory processes underpin morphological variation in these seabirds. More importantly, we detect osteological differentiation between New Zealand and Australian specimens, providing further support for the taxonomic separation of these behaviourally and genetically distinct lineages. Broadly, our study shows the value of multivariate osteological analyses in helping to resolve the status of genetic lineages that appear morphologically ‘cryptic’ when considering external morphometrics and plumage features ...

New Zealand, long recognised as a land where birds dominate the terrestrial vertebrate biota, lacked an informative fossil record for the non-marine pre-Pleistocene avifauna until the twenty-first century. Here we review recent research that alters the known diversity of the fossil Paleogene–Neogene birds and our understanding of the origin of New Zealand’s recent or modern biota. Since 2010, there has been a 50% increase in the number of described fossil bird species (now 45) for the pre-Quaternary period. Many represent higher taxa that are new or listed for New Zealand for the first time, including 12 genera (35 total), nine family-level taxa (18 total), and seven ordinal taxa. We also review recent multidisciplinary research integrating DNA and morphological analyses affecting the taxonomic diversity of the Quaternary avifauna and present revised diversity metrics. The Holocene avifauna contained 217 indigenous breeding species (67% endemic) of which 54 (25%) are extinct.

The relationships of the extinct New Zealand ravens (Corvus spp.) are poorly understood. We sequenced the mitogenomes of the two currently recognised species and found they were sister-taxa to a clade comprising the Australian raven, little raven, and forest raven (C.coronoides, C. mellori and C. tasmanicus respectively). The divergence between the New Zealand ravens and Australian raven clade occurred in the latest Pliocene, which coincides with the onset of glacial deforestation. We also found that the divergence between the two putative New Zealand species C. antipodum and C. moriorum probably occurred in the late Pleistocene making their separation as species untenable. Consequently, we consider Corax antipodum (Forbes, 1893) to be a subspecies of Corvus moriorum Forbes, 1892. We re-examine the osteological evidence that led 19th century researchers to assign the New Zealand taxa to a separate genus, and re-assess these features in light of our new phylogenetic hypotheses. Like previous researchers, we conclude that the morphology of the palate of C. moriorum is unique among the genus Corvus, and suggest this may be an adaptation for a specialist diet.

Museum specimens provide a record of past species distribution and are an increasingly important resource for conservation genetic research. The scientific value of these specimens depends upon the veracity of their associated data and can be compromised by inaccurate details; including taxonomic identity, collection locality, and collector. New Zealand contains many endemic species that have been driven to extinction or reduced to relict distributions following the arrival of humans and mammalian predators, including the Acanthisittid wrens (of which only two of the eight described species presently persist). One of these is the New Zealand rock wren (Xenicus gilviventris), currently classified as an endangered species and experiencing ongoing population declines. Here we analyze ancient DNA retrieved from New Zealand rock wren museum skins to establish the veracity of their recorded collection localities—New Zealand rock wrens exhibit strong north-south genetic structuring along the Southern Alps of New Zealand's South Island. We include the only specimen reportedly collected from New Zealand's North Island, outside the known range of New Zealand rock wrens, specimens collected by Henry Hamersley Travers, a collector known for poor record keeping and potentially fraudulent specimen data, and type specimens of proposed Xenicus taxa. Multiple instances of inaccurate collection locality were detected, including that of the New Zealand rock wren reportedly collected from the North Island, which matches individuals from the southern South Island. Syntypes of X. haasti, and a syntype of X. gilviventris clustered with individuals belonging to the northern New Zealand rock wren lineage. Our results suggest that New Zealand rock wrens have not been historically extirpated from New Zealand's North Island, and that caution must be taken when utilizing museum specimens to inform conservation management decisions. Additionally, we describe the type locality of both X. gilviventris and X. haasti, with genetic and historical evidence suggesting that the specimens used to describe these taxa were collected from the headwaters of the Rakaia River. This study demonstrates that ancient DNA analysis can add value to museum specimens by revealing incorrect specimen data and inform the conservation management and taxonomy of endangered species.

Genetic data are increasingly being used to prioritize species conservation in a fiscally constrained age of seemingly boundless conservation crises. Such data can also reveal previously cryptic biodiversity requiring further revision of conservation management guidelines. Using a combination of mitochondrial (control region) and nuclear (beta fibrinogen intron 7) DNA, and morphology, we reveal that the endemic New Zealand Spotted Shag (Phalacrocorax punctatus) complex exhibits phylogenetic structure that is decoupled from previously recorded qualitative morphological variation. Crucially, the most genetically distinct populations within P. punctatus are from northern New Zealand; recent surveys show that these populations, which house important genetic diversity within Spotted Shags, are in danger of being extirpated. In contrast, we find the previously phenotypically differentiated nominate (P. punctatus punctatus) and Blue (P. punctatus oliveri) Shag subspecies show no genetic and morphological separation, and are of least conservation concern.

Insular avifaunas have repeatedly spawned evolutionary novelties in the form of unusually large, often flightless species. We report fossils from the Early Miocene St Bathans Fauna of New Zealand that attests to the former existence of a giant psittaciform, which is described as a new genus and species. The fossils are two incomplete tibiotarsi from a bird with an estimated mass of 7 kg, double that of the heaviest known parrot, the kakapo Strigops habroptila . These psittaciform fossils show that parrots join the growing group of avian taxa prone to giantism in insular species, currently restricted to palaeognaths, anatids, sylviornithids, columbids, aptornithids, ciconiids, tytonids, falconids and accipitrids.

New Zealand’s unique biodiversity has long been considered both the product of a ‘Moa’s Ark’ isolationist experiment since the break-up of Gondwana and the result of millions of years of transoceanic migration. Increasingly, palaeontological and genetic evidence suggest that much of New Zealand avifauna results from dispersal from Australia. We synthesise a growing number of evolutionary genetic studies to show a previously unrecognised clustering of divergence times in Australian and New Zealand bird species pairs, across the avian phylogeny, around 2.5 million years ago at the beginning of the Pleistocene. The timing coincides with the dramatic environmental changes associated with the Plio-Pleistocene transition. Intriguingly, recent anthropogenic impacts and environmental modifications are replicating in some important ways conditions during the Pleistocene glacial periods, resulting in a new wave of avian ‘native invaders’ into New Zealand.

Background DNA barcoding utilises a standardised region of the cytochrome c oxidase I (COI) gene to identify specimens to the species level. It has proven to be an effective tool for identification of avian samples. The unique island avifauna of New Zealand is taxonomically and evolutionarily distinct. We analysed COI sequence data in order to determine if DNA barcoding could accurately identify New Zealand birds. Results We sequenced 928 specimens from 180 species. Additional Genbank sequences expanded the dataset to 1416 sequences from 211 of the estimated 236 New Zealand species. Furthermore, to improve the assessment of genetic variation in non-endemic species, and to assess the overall accuracy of our approach, sequences from 404 specimens collected outside of New Zealand were also included in our analyses. Of the 191 species represented by multiple sequences, 88.5% could be successfully identified by their DNA barcodes. This is likely a conservative estimate of the power of DNA barcoding in New Zealand, given our extensive geographic sampling. The majority of the 13 groups that could not be distinguished contain recently diverged taxa, indicating incomplete lineage sorting and in some cases hybridisation. In contrast, 16 species showed evidence of distinct intra-species lineages, some of these corresponding to recognised subspecies. For species identification purposes a character-based method was more successful than distance and phylogenetic tree-based methods. Conclusions DNA barcodes accurately identify most New Zealand bird species. However, low levels of COI sequence divergence in some recently diverged taxa limit the identification power of DNA barcoding. A small number of currently recognised species would benefit from further systematic investigations. The reference database and analysis presented will provide valuable insights into the evolution, systematics and conservation of New Zealand birds. Electronic supplementary material The online version of this article (10.1186/s12862-019-1346-y) contains supplementary material, which is available to authorized users.

The recently extinct New Zealand adzebills (Aptornithidae, Aptornis spp.) were an enigmatic group of large flightless birds that have long eluded precise taxonomic assignment as they do not closely resemble any extant birds. Adzebills were nearly wingless, weighed approximately 16&ndash, 19 kg, and possessed massive adze-like reinforced bills whose function remains unknown. Using hybridisation enrichment and high-throughput sequencing of DNA extracted from subfossil bone and eggshell, near-complete mitochondrial genomes were successfully assembled from the two Quaternary adzebill species: the North Island Adzebill (Aptornis otidiformis) and South Island Adzebill (A. defossor). Molecular phylogenetic analyses confirm that adzebills are members of the Ralloidea (rails and allies) and are sister-taxon to the Sarothruridae, which our results suggest comprises the Madagascan wood rails (Mentocrex, two likely sp.) in addition to the tiny (&lt, 50 gram) rail-like Afro-Madagascan flufftails (Sarothrura, 9 spp.). Node age estimates indicate that the split between adzebills and Sarothruridae occurred ~39.6 Ma, suggesting that the ancestors of the adzebills arrived in New Zealand by long-distance dispersal rather than continental vicariance. This newly identified biogeographic link between physically distant New Zealand and Afro-Madagascar, echoed by the relationship between the New Zealand kiwi (Apterygiformes) and Madagascan elephant-birds (Aepyornithiformes), suggests that the adzebill&rsquo, s near relatives were formerly more widespread. In addition, our estimate for the divergence time between the two Quaternary adzebill species (0.2&ndash, 2.3 Ma) coincides with the emergence of a land-bridge between the North and South islands of New Zealand (ca. 1.5&ndash, 2 Ma). This relatively recent divergence suggests that North Island adzebills are the result of a relatively recent dispersal from the South Island, from which the earliest (Miocene) adzebill fossil has been described.

Details of the locality and collection data for the type material of Heracles inexpectatus.

The New Zealand acanthisittid wrens are the sister-taxon to all other "perching birds" (Passeriformes) and - including recently extinct species - represent the most diverse endemic passerine family in New Zealand. Consequently, they are important for understanding both the early evolution of Passeriformes and the New Zealand biota. However, five of the seven species have become extinct since the arrival of humans in New Zealand, complicating evolutionary analyses. The results of morphological analyses have been largely equivocal, and no comprehensive genetic analysis of Acanthisittidae has been undertaken. We present novel mitochondrial genome sequences from four acanthisittid species (three extinct, one extant), allowing us to resolve the phylogeny and revise the taxonomy of acanthisittids. Reanalysis of morphological data in light of our genetic results confirms a close relationship between the extant rifleman (Acanthisitta chloris) and an extinct Miocene wren (Kuiornis indicator), making Kuiornis a useful calibration point for molecular dating of passerines. Our molecular dating analyses reveal that the stout-legged wrens (Pachyplichas) diverged relatively recently from a more gracile (Xenicus-like) ancestor. Further, our results suggest a possible Early Oligocene origin of the basal Lyall's wren (Traversia) lineage, which would imply that Acanthisittidae survived the Oligocene marine inundation of New Zealand and therefore that the inundation was not complete.

Leucocarbo shags are a species-rich seabird clade exhibiting a southern circumpolar distribution. New Zealand's endemic Stewart Island shag, Leucocarbo chalconotus (G. R. Gray, 1845), comprises two regional groups (Otago and Foveaux Strait) that show consistent differences in relative frequencies between pied (black and white) and bronze (wholly dark) plumages, the extent and colour of facial carunculation, body size (based on postcranial morphometrics), and breeding season. Moreover, previous genetic research on modern and historical specimens utilizing mitochondrial DNA control-region sequences has also shown that the Otago and Foveaux lineages may not be sister taxa; instead, in several analyses the Otago lineage is sister to the endemic Chatham Island shag, Leucocarbo onslowi (Forbes, 1893). We present new ancient DNA analyses of the type specimens for the Otago and Foveaux Strait lineages of L. chalconotus, including a phylogenetic reanalysis of the available ancient, historical, and modern control-region sequence data for these lineages (including L. onslowi), and additional statistical analyses incorporating new morphometric characters. These analyses indicate that under the diagnosable species concept the two lineages of Stewart Island shag represent two separate species, which we now recognize as the Otago shag, L. chalconotus (G. R. Gray, 1845), and the Foveaux shag, Leucocarbo stewarti (Ogilvie-Grant, 1898).

Living true seals (phocids) are the most widely dispersed semi-aquatic marine mammals, and comprise geographically separate northern (phocine) and southern (monachine) groups. Both are thought to have evolved in the North Atlantic, with only two monachine lineages—elephant seals and lobodontins—subsequently crossing the equator. The third and most basal monachine tribe, the monk seals, have hitherto been interpreted as exclusively northern and (sub)tropical throughout their entire history. Here, we describe a new species of extinct monk seal from the Pliocene of New Zealand, the first of its kind from the Southern Hemisphere, based on one of the best-preserved and richest samples of seal fossils worldwide. This unanticipated discovery reveals that all three monachine tribes once coexisted south of the equator, and forces a profound revision of their evolutionary history: rather than primarily diversifying in the North Atlantic, monachines largely evolved in the Southern Hemisphere, and from this southern cradle later reinvaded the north. Our results suggest that true seals crossed the equator over eight times in their history. Overall, they more than double the age of the north–south dichotomy characterizing living true seals and confirms a surprisingly recent major change in southern phocid diversity.

We describe a partial skeleton of a stem group penguin from the Waipara Greensand in New Zealand, which is tentatively assigned to Muriwaimanu tuatahi. The fossil includes the first complete wing of a Paleocene penguin and informs on previously unknown features of the mandible and tibiotarsus of small-sized Sphenisciformes from the Waipara Greensand. The wing is distinguished by important features from that of all geologically younger Sphenisciformes and documents an early stage in the evolution of wing-propelled diving in penguins. In particular, the wing of the new fossil exhibits a well-developed alular phalanx and the distal phalanges are not flattened. Because the wing phalanges resemble those of volant birds, we consider it likely that the wing feathers remained differentiated into functional categories and were not short and scale-like as they are in extant penguins. Even though the flippers of geologically younger penguins may favor survival in extremely cold climates, they are likely to have been shaped by hydrodynamic demands. Possible selective drivers include a diminished importance of the hindlimbs in subaquatic propulsion, new foraging strategies (the caudal end of the mandible of the new fossil distinctly differs from that of extant penguins), or increased predation by marine mammals.