Alan J. D. Tennyson

A working list of breeding bird species of the New Zealand region at first human contact

Abstract We present an annotated working list of the bird species breeding in New Zealand during the late Pleistocene and Holocene, up to the time of human contact. New Zealand is defined as including the three main islands and the surrounding smaller islands, plus outlying island groups from Norfolk Island in the northwest, the Kermadec, Chatham, Bounty, Antipodes, Campbell, Auckland, Snares, to Macquarie Islands, but excluding islands south of Macquarie Island and the Ross Dependency. Inclusions or exclusions of species from the list were based on specified criteria. We include only species with a breeding population and not vagrants that occur in New Zealand but which breed elsewhere. Species with validly published names were included if there was fossil evidence for a breeding population before human contact. Species with a breeding population at the time of European contact were included unless contrary evidence from the fossil record indicates that they actually colonised after human settlement. Species without a fossil record were included if a breeding population exists on a relatively undisturbed island within the New Zealand archipelago as defined above. Species now present on the main islands were excluded if they are absent from all well‐documented fossil faunas. Species were excluded from the breeding fauna and treated as vagrants where sustained breeding has not been demonstrated. The phylogenetic species concept is applied both to fossil and to living taxa. The late Quaternary fossil record of birds in New Zealand is excellent, and the contribution of extinct taxa to the total list is understood at least as well as that of the surviving taxa. Many taxa presently recognised at subspecific level are treated here as full species. Twelve extinct species whose former presence is known from fossil evidence, but for which no description has been published, are listed under informal species designations. Taxonomic considerations limited the extent to which the main list could reflect present understanding of the diversity of the avifauna; some undescribed species are at present subsumed under one species name. Where previous taxonomic publications provide precedence, available names at the species‐level have been used. A supplementary hypothetical species list includes all nomenclatural changes signalled in extensive annotations to the main list. In this list we accept 245 species in 110 genera representing 46 families; 176 species were endemic to the archipelago. Preliminary biogeographic analyses based on the composition of the supplementary list show that there were four separate regional faunas: a northern subtropical fauna (Norfolk, Kermadecs); the major fauna of the main islands (North, South, Stewart, and offshore islands); a Chathams fauna (Chatham Islands only); and a subantarctic fauna on the southern islands. Species with wider distributions formed link groups. The origin and compositions of the regional avifaunas and their endemic species differ with their geographic position, climate, and proximity to source faunas. Instances of speciation in groups such as the Coenocoiypha snipe and Petroica flycatchers, and adaptive radiations in groups including moa and acanthisittid wrens, show that there are many avenues for research on the rate of evolution in island and mainland populations of New Zealand birds and that there are large gaps in knowledge of even common taxa. A brief case study demonstrates the inadequacies of using species lists that do not include Holocene fossil species. Species‐area curves based on the total fauna differ substantially from those developed in previous studies based on incomplete, or biased, lists. Pleistocene glaciations caused the pattern of distribution of species on the main islands to change in concert with vegetation changes. Other possible effects include the elimination of warm climate species early in the cooling phase more than 1 million years ago, the speciation in groups including waders and parrots as new habitats (e.g., braided riverbeds and alpine areas) appeared, and the appearance regularly during the Pleistocene of islands that were potential staging points for colonisation of the Chatham Islands. For at least the past 100 000 years, until 2000 years ago, the fauna appears to have been very stable in composition, despite strong cyclic fluctuations in climate and vegetation. The effects of extinctions within the past 2000 years on the composition of the present fauna include the elimination of most of the endemic taxa from all but the subantarctic faunas. Only 169 species of the original late Holocene breeding fauna survive. The extinctions have resulted in a strong bias towards marine and coastal taxa in the present avifauna, in contrast to the balanced representation of terrestrial and marine species in the Pleistocene and Holocene fauna. The importance of systematic studies and the determination of the status of island populations to conservation and basic ornithological research is emphasised. The systematic status of many New Zealand birds is poorly known at present.

Miocene mammal reveals a Mesozoic ghost lineage on insular New Zealand, southwest Pacific

New Zealand (NZ) has long been upheld as the archetypical example of a land where the biota evolved without nonvolant terrestrial mammals. Their absence before human arrival is mysterious, because NZ was still attached to East Antarctica in the Early Cretaceous when a variety of terrestrial mammals occupied the adjacent Australian portion of Gondwana. Here we report discovery of a nonvolant mammal from Miocene (19–16 Ma) sediments of the Manuherikia Group near St Bathans (SB) in Central Otago, South Island, NZ. A partial relatively plesiomorphic femur and two autapomorphically specialized partial mandibles represent at least one mouse-sized mammal of unknown relationships. The material implies the existence of one or more ghost lineages, at least one of which (based on the relatively plesiomorphic partial femur) spanned the Middle Miocene to at least the Early Cretaceous, probably before the time of divergence of marsupials and placentals >125 Ma. Its presence in NZ in the Middle Miocene and apparent absence from Australia and other adjacent landmasses at this time appear to reflect a Gondwanan vicariant event and imply persistence of emergent land during the Oligocene marine transgression of NZ. Nonvolant terrestrial mammals disappeared from NZ some time since the Middle Miocene, possibly because of late Neogene climatic cooling.

Miocene waterfowl and other birds from central Otago, New Zealand

Synopsis Abundant fossil bird bones from the lower Bannockburn Formation, Manuherikia Group, an Early‐Middle Miocene lacustrine deposit, 16–19 Ma, from Otago in New Zealand, reveal the “St Bathans Fauna” (new name), a first Tertiary avifauna of land and freshwater birds from New Zealand. At least 23 species of birds are represented by bones, and probable moa, Aves: Dinornithiformes, by eggshell. Anatids dominate the fauna with four genera and five species described as new: a sixth and largest anatid species is represented by just one bone. This is the most diverse Early‐Middle Miocene duck fauna known worldwide. Among ducks, two species of dendrochenines are most numerous in the fauna, but a tadornine is common as well. A diving petrel (Pelecanoididae: Pelecanoides) is described, so extending the geological range of this genus worldwide from the Pliocene to the Middle Miocene, at least. The remaining 16 taxa are left undescribed but include: a large species of gull (Laridae); two small waders (Charadriiformes, genus indet.), the size of Charadrius bicinctus and Calidris ruficollis, respectively; a gruiform represented by one specimen similar to Aptornis; abundant rail (Rallidae) bones, including a common flightless rail and a rarer slightly larger taxon, about the size of Gallirallus philippensis; an ?eagle (Accipitridae); a pigeon (Columbidae); three parrots (Psittacidae); an owlet nightjar (Aegothelidae: Aegotheles sp.); a swiftlet (Apodidae: Collocalia sp.); and three passerine taxa, of which the largest is a member of the Cracticidae. The absence of some waterbirds, such as anserines (including swans), grebes (Podicipedidae) and shags (Phalacrocoracidae), among the abundant bones, indicates their probable absence from New Zealand in the Early‐Middle Miocene.

A sphenodontine (Rhynchocephalia) from the Miocene of New Zealand and palaeobiogeography of the tuatara (Sphenodon)

Jaws and dentition closely resembling those of the extant tuatara (Sphenodon) are described from the Manuherikia Group (Early Miocene; 19–16 million years ago, Mya) of Central Otago, New Zealand. This material is significant in bridging a gap of nearly 70 million years in the rhynchocephalian fossil record between the Late Pleistocene of New Zealand and the Late Cretaceous of Argentina. It provides the first pre-Pleistocene record of Rhynchocephalia in New Zealand, a finding consistent with the view that the ancestors of Sphenodon have been on the landmass since it separated from the rest of Gondwana 82–60 Mya. However, if New Zealand was completely submerged near the Oligo-Miocene boundary (25–22 Mya), as recently suggested, an ancestral sphenodontine would need to have colonized the re-emergent landmass via ocean rafting from a currently unrecorded and now extinct Miocene population. Although an Early Miocene record does not preclude that possibility, it substantially reduces the temporal window of opportunity. Irrespective of pre-Miocene biogeographic history, this material also provides the first direct evidence that the ancestors of the tuatara, an animal often perceived as unsophisticated, survived in New Zealand despite substantial local climatic and environmental changes.

Biogeographical and phylogenetic implications of an early Miocene wren (Aves: Passeriformes: Acanthisittidae) from New Zealand

ABSTRACT A new species and genus of acanthisittid wren (Aves: Passeriformes: Acanthisittidae) is described from the Early Miocene (19–16 Ma) St Bathans Fauna from Otago, New Zealand, based on four fossil bones. The first Tertiary fossil passerine to be described from New Zealand, it is similar in size to New Zealands smallest extant bird, the Rifleman Acanthisitta chloris. A phylogenetic analysis of 53 osteological characters and 24 terminal taxa, including four suboscines, basal corvoids (Menuridae, Atrichornithidae, Climacteridae, Ptilonorhynchidae, Maluridae, Dasyornithidae, Acanthizidae, Pardalotidae, Meliphagidae), and all seven Recent acanthisittid species, identifies it as the sister group to Acanthisitta. This, the first phylogenetic analysis of the basal passerine groups to use morphological characters, recovers a similar pattern of relationships of basal corvoid taxa to that obtained by recent molecular studies. The analysis also suggests that Acanthisitta chloris and the new species are the most deeply nested taxa within the family, suggesting that the radiation of Recent acanthisittids originated no later than the Early Miocene.

Miocene skinks and geckos reveal long-term conservatism of New Zealand's lizard fauna

The New Zealand (NZ) lizard fossil record is currently limited to late Quaternary remains of modern taxa. The St Bathans Fauna (early Miocene, southern South Island) extends this record to 19–16 million years ago (Myr ago). Skull and postcranial elements are similar to extant Oligosoma (Lygosominae) skinks and Hoplodactylus (Diplodactylinae) geckos. There is no evidence of other squamate groups. These fossils, along with coeval sphenodontines, demonstrate a long conservative history for the NZ lepidosaurian fauna, provide new molecular clock calibrations and contradict inferences of a very recent (less than 8 Myr ago) arrival of skinks in NZ.

Bone growth marks reveal protracted growth in New Zealand kiwi (Aves, Apterygidae)

The presence of bone growth marks reflecting annual rhythms in the cortical bone of non-avian tetrapods is now established as a general phenomenon. In contrast, ornithurines (the theropod group including modern birds and their closest relatives) usually grow rapidly in less than a year, such that no annual rhythms are expressed in bone cortices, except scarce growth marks restricted to the outer cortical layer. So far, cyclical growth in modern birds has been restricted to the Eocene Diatryma, the extant parrot Amazona amazonica and the extinct New Zealand (NZ) moa (Dinornithidae). Here we show the presence of lines of arrested growth in the long bones of the living NZ kiwi (Apteryx spp., Apterygidae). Kiwis take 5–6 years to reach full adult body size, which indicates a delayed maturity and a slow reproductive cycle. Protracted growth probably evolved convergently in moa and kiwi sometime since the Middle Miocene, owing to the severe climatic cooling in the southwest Pacific and the absence of mammalian predators.

A Large Fruit Pigeon (Columbidae) from the Early Miocene of New Zealand

ABSTRACT. We describe a new genus and species of pigeon (Columbiformes) from a single coracoid from the St Bathans Fauna of New Zealand (16–19 mya). It is the first columbid species described from pre-Pliocene deposits in Australasia. Two apomorphies identify the fossil as belonging to the ptilinopine group of fruit pigeons, among which it is most similar to Hemiphaga, the large fruit pigeon currently endemic to the New Zealand biogeographic area. This reveals that the Hemiphaga lineage has been in New Zealand since the Early Miocene, which supports recent divergence-date estimates for Hemiphaga and its modern sister taxon (Lopholaimus) based on molecular data.

Fish Remains, Mostly Otoliths, from the Non-Marine Early Miocene of Otago, New Zealand

Fish remains described from the early Miocene lacustrine Bannockburn Formation of Central Otago, New Zealand, consist of several thousand otoliths and one skeleton plus another disintegrated skull. One species, Mataichthys bictenatus Schwarzhans, Scofield, Tennyson, and T. Worthy gen. et sp. nov., an eleotrid, is established on a skeleton with otoliths in situ. The soft embedding rock and delicate, three-dimensionally preserved fish bones were studied by CT-scanning technology rather than physical preparation, except where needed to extract the otolith. Fourteen species of fishes are described, 12 new to science and two in open nomenclature, representing the families Galaxiidae (Galaxias angustiventris, G. bobmcdowalli, G. brevicauda, G. papilionis, G. parvirostris, G. tabidus), Retropinnidae (Prototroctes modestus, P. vertex), and Eleotridae (Mataichthys bictenatus, M. procerus, M. rhinoceros, M. taurinus). These findings prove that most of the current endemic New Zealand/southern Australia freshwater fish fauna was firmly established in New Zealand as early as 19–16 Ma ago. Most fish species indicate the presence of large fishes, in some cases larger than Recent species of related taxa, for instance in the eleotrid genus Mataichthys when compared to the extant Gobiomorphus. The finding of a few otoliths from marine fishes corroborates the age determination of the Bannockburn Formation as the Altonian stage of the New Zealand marine Tertiary stratigraphy.

An Early Miocene Diversity of Parrots (Aves, Strigopidae, Nestorinae) from New Zealand

ABSTRACT A new genus and three species of parrot (Psittaciformes, Strigopidae, Nestorinae) are described from the early Miocene (19–16 Ma) St Bathans Fauna of Otago, New Zealand, based on 85 fossils as follows: Nelepsittacus minimus (17), N. donmertoni (60), and N. daphneleeae (6), with two additional fossils representing a fourth unnamed taxon. These taxa range from small parrots approximately the size of Cyanoramphus species to one as large as the living Nestor notabilis. Apomorphies in the coracoid, humerus, tibiotarsus, and tarsometatarsus ally Nelepsittacus with Nestor and exclude a close relationship with Strigops, the other endemic genus assumed to have had a long history in New Zealand. With only nestorine parrots represented, the St Bathans Fauna has nothing in common with the Australian psittaciform fauna, in which cacatuids and a diversity of psittacid genera exist. These data add to the growing body of evidence that the New Zealand terrestrial vertebrate fauna, at a time minimally 3 Ma after the maximal marine inundation of Zealandia in the late Oligocene, was highly endemic, with no close relationship to the closest faunas in Australia. This high degree of endemism strongly suggests that the Zealandian terrestrial biota persisted, at least in part, through the Oligocene highstand in sea level.