Dinosaur sanctuary on the Chatham Islands, Southwest Pacific: First record of theropods from the K–T boundary Takatika Grit

Cretaceous–Tertiary (K–T) boundary (ca. 65 Ma) sections on a Southwest Pacific island containing dinosaurs were unknown until March 2003 when theropod bones were recovered from the Takatika Grit on the remote Chatham Islands (latitude 44° S, longitude 176° W), along the Chatham Rise. Tectonic and palaeontologic evidence support the eastward extension of a ca. 900 km land bridge that connected the islands to what is now New Zealand prior to the K–T boundary. The Chathams terrestrial fauna inhabited coastal, temperate environments along a low-lying, narrow, crustal extension of the New Zealand subcontinent, characterised by a tectonically dynamic, volcanic landscape with eroding hills (horsts) adjacent to flood plains and deltas, all sediments accumulating in grabens. This finger-like tract was blanketed with a conifer and clubmoss (Lycopodiopsida) dominated forest. The Chatham Islands region would have, along with New Zealand, provided a dinosaur island sanctuary after separating from the Gondwana margin ca. 80 Ma.     

Multiple colonizations of a remote oceanic archipelago by one species: how common is long‐distance dispersal?

Aim  It is well established that many groups of plants and animals have undergone long-distance dispersal, but the extent to which this continues beyond initial colonization is largely unknown. To provide further insight into the frequency of gene flow mediated by long-distance dispersal, we investigated the origins of the fern Asplenium hookerianum on the Chatham Islands, and present a review of the contribution of molecular data to elucidating the origins of this archipelago's biota.
Location  Chatham Islands and New Zealand. A. hookerianum is scarce on the Chatham Islands but common in New Zealand, some 800 km to the west.
Methods  We compared chloroplast trnL–trnF DNA sequence data from Chatham Islands' A. hookerianum with extensive phylogeographic data for this genetically variable species in mainland New Zealand.
Results  Our sequencing revealed the presence of two haplotypes in Chatham Islands' A. hookerianum . These haplotypes differed by four mutational events and were each more closely related to haplotypes found in New Zealand than to each other.
Main conclusions  Despite the rarity of A. hookerianum on the Chatham Islands, its populations there appear to derive from at least two long-distance dispersal events from New Zealand, these possibly originating from different areas. We suggest that long-distance transoceanic dispersal, and the gene flow it can mediate, may be more common than is generally appreciated.     

Pilot trophic model for subantarctic water over the Southern Plateau, New Zealand: a low biomass, high transfer efficiency system

The Southern Plateau subantarctic region, southeast of New Zealand, is an important feeding area for birds, seals and fish, and a fishing ground for commercially significant species. The Southern Plateau is a major morphometric feature, covering approximately 433,620 km² with average depth of 615 m. The region is noted for its relatively low levels of phytoplankton biomass and primary production that is iron-limited. In order to evaluate the implications of these attributes for the functioning of this ecosystem a steady-state, 19-compartment model was constructed using Ecopath with Ecosim software of Christensen et al. [www.ecopath.org]. The system is driven by primary production that is primarily governed by the supply of iron and light. The total system biomass of 6.28 g C m⁻² is very low compared with systems so far modelled with a total system throughput of 1136 g C m⁻² year⁻¹. In the model, the Southern Plateau retains 69% of the biomass in the pelagic system and 99% of total production. Although fish are caught demersally, most of their food is part of production in the pelagic system. Top predators represent about 0.3% of total biomass and account for about 0.24 g C m⁻² year⁻¹ of food consumed made up of birds 0.058 g C m⁻² year⁻¹, seals 0.041 g C m⁻² year⁻¹, and toothed 0.094 g C m⁻² year⁻¹ and baleen whales 0.051 g C m⁻² year⁻¹. This amounts to 105,803 tonnes carbon over the whole of the Southern Plateau and is about 17% of the total amount of food eaten by non-mesopelagic fish. Mean transfer efficiencies between trophic levels II and IV of 23% are at the high end of the range reported in the literature. In the model, adult fish production is almost completely accounted for by the fisheries take (32%), consumption by seals (7%), toothed whales (21%), other adult fish (13%), and squid (20%). Fish and squid catches are at the trophic levels of 4.8 and 5.0, respectively. The gross efficiency of the fishery is 0.018% (catch/primary production). Although not all data come from direct knowledge of this system, the model reflects its general characteristics, namely a low primary production system dominated by the microbial loop, low sedimentation to the seafloor, high transfer efficiencies, a long food web and supporting high-level predators.     

Population structure and biogeography of Hemiphaga pigeons (Aves: Columbidae) on islands in the New Zealand region

Aim The New Zealand avifauna includes lineages that lack close relatives elsewhere and have low diversity, characteristics sometimes ascribed to long geographic isolation. However, extinction at the population and species levels could yield the same pattern. A prominent example is the ecologically important pigeon genus Hemiphaga. In this study, we examined the population structure and phylogeography of Hemiphaga across islands in the region. Location New Zealand, Chatham Islands and Norfolk Island. Methods Mitochondrial DNA was sequenced for all species of the genus Hemiphaga. Sixty‐seven individuals from mainland New Zealand (Hemiphaga novaeseelandiae novaeseelandiae), six of the Chatham Islands sister species (Hemiphaga chathamensis), and three of the extinct Norfolk Island subspecies (Hemiphaga novaeseelandiae spadicea) were included in this study. Novel D‐loop and cytochrome b primers were designed to amplify DNA from museum samples. Additionally, five other mitochondrial genes were used to examine placement of the phylogenetic root. Results Analyses of mitochondrial DNA sequences revealed three Hemiphaga clades, consistent with the allopatric populations of recognized (sub)species on oceanic islands. Of the 23 D‐loop haplotypes among 67 New Zealand pigeons (Hemiphaga n. novaeseelandiae), 19 haplotypes were singletons and one haplotype was common and widespread. Population genetic diversity was shallow within and between New Zealand populations, indicating range expansion with high inter‐population exchange. Tentative rooting of the Hemiphaga clade with cyt b data indicates exchange between mainland New Zealand and the Chatham Islands prior to colonization of Norfolk Island. We found low genetic divergence between populations on New Zealand, the Chatham Islands and Norfolk Island, but deep phylogenetic divergence from the closest living relatives of Hemiphaga. Main conclusions The data are consistent with the hypothesis of population reduction during the Pleistocene and subsequent expansion from forest refugia. Observed mobility of Hemiphaga when feeding helps explain the shallow diversity among populations on islands separated by many hundreds of kilometres of ocean. Together with comparison of distribution patterns observed among birds of the New Zealand region, these data suggest that endemicity might represent not long occupancy of an area, but descent from geologically recent colonizations. We consider the role of lineage pruning in creating the impression of old endemicity.     

Panbiogeography of New Caledonia,south‐west Pacific: basal angiosperms on basement terranes,ultramafic endemics inherited from volcanic island arcs and old taxa endemic to young islands

Aim To investigate areas of endemism in New Caledonia and their relationship with tectonic history. Location New Caledonia, south‐west Pacific. Methods Panbiogeographical analysis. Results Biogeographical patterns within New Caledonia are described and illustrated with reference to eight terranes and ten centres of endemism. The basement terranes make up a centre of endemism for taxa including Amborella, the basal angiosperm. Three of the terranes that accreted to the basement in the Eocene (high‐pressure metamorphic terrane, ultramafic nappe and Loyalty Ridge) have their own endemics. Main conclusions New Caledonia is not simply a fragment of Gondwana but, like New Zealand and New Guinea, is a complex mosaic of allochthonous terranes. The four New Caledonian basement terranes were all formed from island arc‐derived and arc‐associated material (including ophiolites) which accumulated in the pre‐Pacific Ocean, not in Gondwana. They amalgamated and were accreted to Gondwana (eastern Australia) in the Late Jurassic/Early Cretaceous, but in the Late Cretaceous they separated from Australia with the opening of the Tasman Sea and break‐up of Gondwana. An Eocene collision of the basement terranes with an island arc to the north‐east – possibly the Loyalty Ridge – is of special biogeographical interest in connection with New Caledonia–central Pacific affinities. The Loyalty–Three Kings Ridge has had a separate history from that of the Norfolk Ridge/New Caledonia, although both now run in parallel between Vanuatu and New Zealand. The South Loyalty Basin opened between Grande Terre and the Loyalty Ridge in the Cretaceous and attained a width of 750 km. However, it was almost completely destroyed by subduction in the Eocene which brought the Loyalty Ridge and Grande Terre together again, after 30 Myr of separation. The tectonic history is reflected in the strong biogeographical differences between Grande Terre and the Loyalty Islands. Many Loyalty Islands taxa are widespread in the Pacific but do not occur on Grande Terre, and many Grande Terre/Australian groups are not on the Loyalty Islands. The Loyalty Islands are young (2 Myr old) but they are merely the currently emergent parts of the Loyalty Ridge whose ancestor arcs have a history of volcanism dating back to the Cretaceous. Old taxa endemic to the young Loyalty Ridge islands persist over geological time as a dynamic metapopulation surviving in situ on the individually ephemeral islands and atolls found around subduction zones. The current Loyalty Islands, like the Grande Terre terranes, have inherited their biota from previous islands. On Grande Terre, the ultramafic terrane was emplaced on Grande Terre in the Eocene (about the same time as the collision with the island arc). The very diverse endemic flora on the ultramafics may have been inherited by the obducting nappe from prior base‐rich habitat in the region, including the mafic Poya terrane and the limestones typical of arc and intraplate volcanic islands.     

From East Gondwana to Central America: historical biogeography of the Alstroemeriaceae

Aim The Alstroemeriaceae is among 28 angiosperm families shared between South America, New Zealand and/or Australia; here, we examine the biogeography of Alstroemeriaceae to better understand the climatic and geological settings for its diversification in the Neotropics. We also compare Alstroemeriaceae with the four other Southern Hemisphere families that expanded from Patagonia to the equator, to infer what factors may have permitted such expansions across biomes. Location South America, Central America, Australia and New Zealand. Methods Three chloroplast genes, one mitochondrial gene and one nuclear DNA region were sequenced for 153 accessions representing 125 of the 200 species of Alstroemeriaceae from throughout the distribution range; 25 outgroup taxa were included to securely infer evolutionary directions and be able to use both ingroup and outgroup fossil constraints. A relaxed‐clock model relied on up to three fossil calibrations, and ancestral ranges were inferred using statistical dispersal–vicariance analysis (S‐DIVA). Southern Hemisphere disjunctions in the flowering plants were reviewed for key biological traits, divergence times, migration directions and habitats occupied. Results The obtained chronogram and ancestral area reconstruction imply that the most recent common ancestor of Colchicaceae and Alstroemeriaceae lived in the Late Cretaceous in southern South America/Australasia, the ancestral region of Alstroemeriaceae may have been South America/Antarctica, and a single New Zealand species is due to recent dispersal from South America. Chilean Alstroemeria diversified with the uplift of the Patagonian Andes c. 18 Ma, and a hummingbird‐pollinated clade (Bomarea) reached the northern Andes at 11–13 Ma. The South American Arid Diagonal (SAAD), a belt of arid vegetation caused by the onset of the Andean rain shadow 14–15 Ma, isolated a Brazilian clade of Alstroemeria from a basal Chilean/Argentinean grade. Main conclusions Only Alstroemeriaceae, Calceolariaceae, Cunoniaceae, Escalloniaceae and Proteaceae have expanded and diversified from Patagonia far into tropical latitudes. All migrated northwards along the Andes, but also reached south‐eastern Brazil, in most cases after the origin of the SAAD. Our results from Alstroemeria now suggest that the SAAD may have been a major ecological barrier in southern South America.     

Australian Oligo-Miocene Mystacinids (Microchiroptera): upper dentition, new taxa and divergence of New Zealand species

The upper dentition of two Australian early Miocene mystacinids, Icaropsparadox and I. aenae, from Riversleigh, Queensland, are described for the first time. Also recognised is a late Oligocene mystacinid from Lake Palankarinna, South Australia. The new fossils help refine understanding about the evolutionary history of mystacinids in Australia, including their temporal and geographical range, possible dietary and roosting habits, and likely separation time of New Zealand mystacinids.     

A diverse fossil terrestrial arthropod fauna from New Zealand: evidence from the early Miocene Foulden Maar fossil lagerstätte

Fossil evidence for the evolutionary history of terrestrial arthropods in New Zealand is extremely limited; only six pre‐Quaternary insects (Triassic to Eocene) have been recorded previously, none of Miocene age. The Foulden Maar fossil lagerstätte in Otago has now yielded a diverse arthropod assemblage, including members of the Araneae, Plecoptera, Isoptera, Hemiptera, Coleoptera, Hymenoptera, Trichoptera and Diptera. The fauna significantly emends the fossil record for the Southern Hemisphere, provides an unparalleled insight into a 23‐million‐year‐old New Zealand lake/forest palaeoecosystem and allows a first evaluation of arthropod diversity at a time coeval with or shortly after the maximum marine transgression of Zealandia in the late Oligocene. The well‐preserved arthropods chiefly represent ground‐dwelling taxa of forest floor and leaf litter habitats, mostly from sub‐families and genera that are still present in the modern fauna. They provide precisely dated fossil evidence for the antiquity of some of New Zealand's terrestrial arthropods and the first potential time calibrations for phylogenetic studies. The high arthropod diversity at Foulden Maar, together with a subtropical rainforest flora and fossil evidence for complex arthropod–plant interactions, suggests that terrestrial arthropods persisted during the Oligocene marine transgression of Zealandia.     

Human‐mediated extirpation of the unique Chatham Islands sea lion and implications for the conservation management of remaining New Zealand sea lion populations

While terrestrial megafaunal extinctions have been well characterized worldwide, our understanding of declines in marine megafauna remains limited. Here, we use ancient DNA analyses of prehistoric (<1450–1650 AD) sea lion specimens from New Zealand's isolated Chatham Islands to assess the demographic impacts of human settlement. These data suggest there was a large population of sea lions, unique to the Chatham Islands, at the time of Polynesian settlement. This distinct mitochondrial lineage became rapidly extinct within 200 years due to overhunting, paralleling the extirpation of a similarly large endemic mainland population. Whole mitogenomic analyses confirm substantial intraspecific diversity among prehistoric lineages. Demographic models suggest that even low harvest rates would likely have driven rapid extinction of these lineages. This study indicates that surviving Phocarctos populations are remnants of a once diverse and widespread sea lion assemblage, highlighting dramatic human impacts on endemic marine biodiversity. Our findings also suggest that Phocarctos bycatch in commercial fisheries may contribute to the ongoing population decline.     

Evolutionary and biogeographical history of an ancient and global group of arachnids (Arachnida: Opiliones: Cyphophthalmi) with a new taxonomic arrangement

We investigate the phylogeny, biogeography, time of origin and diversification, ancestral area reconstruction and large‐scale distributional patterns of an ancient group of arachnids, the harvestman suborder Cyphophthalmi. Analysis of molecular and morphological data allow us to propose a new classification system for the group; Pettalidae constitutes the infraorder Scopulophthalmi new clade , sister group to all other families, which are divided into the infraorders Sternophthalmi new clade and Boreophthalmi new clade . Sternophthalmi includes the families Troglosironidae, Ogoveidae, and Neogoveidae; Boreophthalmi includes Stylocellidae and Sironidae, the latter family of questionable monophyly. The internal resolution of each family is discussed and traced back to its geological time origin, as well as to its original landmass, using methods for estimating divergence times and ancestral area reconstruction. The origin of Cyphophthalmi can be traced back to the Carboniferous, whereas the diversification time of most families ranges between the Carboniferous and the Jurassic, with the exception of Troglosironidae, whose current diversity originates in the Cretaceous/Tertiary. Ancestral area reconstruction is ambiguous in most cases. Sternophthalmi is traced back to an ancestral land mass that contained New Caledonia and West Africa in the Permian, whereas the ancestral landmass for Neogoveidae included the south‐eastern USA and West Africa, dating back to the Triassic. For Pettalidae, most results include South Africa, or a combination of South Africa with the Australian plate of New Zealand or Sri Lanka, as the most likely ancestral landmass, back in the Jurassic. Stylocellidae is reconstructed to the Thai‐Malay Penisula during the Jurassic. Combination of the molecular and morphological data results in a hypothesis for all the cyphophthalmid genera, although the limited data available for some taxa represented only in the morphological partition negatively affects the phylogenetic reconstruction by decreasing nodal support in most clades. However, it resolves the position of many monotypic genera not available for molecular analysis, such as Iberosiro, Odontosiro, Speleosiro, Managotria or Marwe, although it does not place Shearogovea or Ankaratra within any existing family. The biogeographical data show a strong correlation between relatedness and formerly adjacent landmasses, and oceanic dispersal does not need to be postulated to explain disjunct distributions, especially when considering the time of divergence. The data also allow testing of the hypotheses of the supposed total submersion of New Zealand and New Caledonia, clearly falsifying submersion of the former, although the data cannot reject the latter. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 92–130.     

A biogeographic review of Parahebe (Scrophulariaceae)

Distribution maps and notes are given for the 41 species of Parahebe sensu lata. The genus occurs in New Zealand, south-east Australia and New Guinea, with greatest diversity in New Zealand, especially in the Spenser Mountains region of South Island. A group of species with ciliate floral discs is found in north-east South Island, and also in eastern Papua New Guinea. This outer Australasian arc distribution is attributed to the group having originated before the break-up of Gondwana. Within New Zealand the P. catarraclae complex shows disjunction along the Alpine Fault, a plate boundary of the transform type. The disjunction is attributed to massive lateral displacement on the Fault during Tertiary time pulling apart plant populations. Parahebe sect. Paniculatae is newly described. The following new combinations are made: Parahebe brevistylis, P. macrantha, P. macrantha var. brachyphylla, P. raoulii, P. r. subsp. maccaskillii, P. r. subsp. pentasepala, P. lavaudiana, P. hulkeana, P. nivea, P. arenaria, P. velutina, P. blakelyi, P. arcuata, P. derwentiana subsp. maideniana, P. d. subsp. homalodonta, P. d. subsp. anisodonta and P. d. subsp. subglauca.     

Biogeographical affinities of the New Caledonian biota: a puzzle with 24 pieces

Aim The distributions of many New Caledonian taxa were reviewed in order to ascertain the main biogeographical connections with other areas. Location Global. Methods Panbiogeographical analysis. Results Twenty‐four areas of endemism (tracks) involving New Caledonia and different areas of Gondwana, Tethys and the central Pacific were retrieved. Most are supported by taxa of lower and higher plants, and lower and higher animals. Main conclusions Although parts of New Caledonia were attached to Gondwana for some time in the mid‐Cretaceous, most of the New Caledonian terranes formed as oceanic island arcs and sections of sea floor bearing seamounts. The flora and fauna have evolved and survived for tens of millions of years as metapopulations on ephemeral islands. Later, the biotas were juxtaposed and fused during terrane accretion. This process, together with the rifting of Gondwana, explains the biogeographical affinities of New Caledonia with parts of Gondwana, Tethys and the Pacific.     

Surface movement rates of the New Zealand flatworm Artioposthia triangulata: potential for spread by active migration

Movement rates for the surface activity of the New Zealand flatworm Artioposthia triangulata (Dendy) (Tricladida: Terricola) were estimated under semi-field conditions in the Faroe Islands. The distance travelled during 10 consecutive 1-minute periods per individual was measured from mucus trails left behind as the flatworms moved over black plastic sheeting sprayed with fine droplets of water. Adult specimens were found to crawl at rates of up to 28 cm min^-1 (almost 17m h^-1), whereas juveniles crawled at rates of between 3 and 4cm min^-1 (approximately 2m h^-1). This study provides the first data concerning the potential for species spread by active migration.     

The evolution of approach herkogamy from protandry in New ZealandGentiana (Gentianaceae)

Most mainland New Zealand species ofGentiana are protandrous as are manyGentianaceae elsewhere in the world. Selfing has evolved in some species confined to areas where pollinator availability is limited by weather, for example, inG. lineata. InG. chathamica, endemic to the Chatham Islands, dichogamy has been lost without the evolution of selfing; instead the flowers are approach herkogamous. It is suggested that as herkogamous blossoms can both receive and dispatch pollen during a single insect visit, whereas dichogamous systems usually require two visits, such a system may have evolved inG. chathamica to maintain outcrossing in an area where prospects for insect pollination are poor.     

陇东黄土高原天然草地植被类型及特征

陇东(庆阳)黄土高原天然草地植被可划分为草甸草原区和典型草原区2个植被区域,包括温性疏灌草丛、草甸草原和典型草原3个植被型14个群系,分别是:酸枣-杠柳-蒿类温性灌草丛、蕤核-红花锦鸡儿-蒿类温性灌草丛、杠柳-蕤核-蒿类-白羊草温性灌草丛、酸枣-蒿类-白羊草温性灌草丛、酸枣-河蒴荛花-白羊草温性灌草丛、白草- 鹅观草-达乌里胡枝子-蒿类暖性灌草丛、白羊草草原、白草草原、赖草草原、蒿类-白羊草草甸草原、长芒草草原、百里香草原、铁杆蒿-茭蒿草原、毛沙芦草草原。     

Evolutionary size changes in plants of the south‐west Pacific

Aim To investigate evolutionary changes in the size of leaves, stems and seeds of plants inhabiting isolated islands surrounding New Zealand. Location Antipodes, Auckland, Campbell, Chatham, Kermadec, Three Kings and Poor Knights Islands. Methods First, we compared the size of leaves and stems produced by 14 pairs of plant taxa between offshore islands and the New Zealand mainland, which were grown in a common garden to control for environmental effects. Similar comparisons of seed sizes were made between eight additional pairs of taxa. Second, we used herbarium specimens from 13 species pairs to investigate scaling relationships between leaf and stem sizes in an attempt to pinpoint which trait might be under selection. Third, we used herbarium specimens from 20 species to test whether changes in leaf size vary among islands located at different latitudes. Lastly, we compiled published records of plant heights to test whether insular species in the genus Hebe differed in size from their respective subgenera on the mainland. Results Although some evidence of dwarfism was observed, most insular taxa were larger than their mainland relatives. Leaf sizes scaled positively with stem diameters, with island taxa consistently producing larger leaves for any given stem size than mainland species. Leaf sizes also increased similarly among islands located at different latitudes. Size changes in insular Hebe species were unrelated to the average size of the respective subgenera on the mainland. Main conclusions Consistent evidence of gigantism was observed, suggesting that plants do not obey the island rule. Because our analyses were restricted to woody plants, results are also inconsistent with the ‘weeds‐to‐trees’ hypothesis. Disproportionate increases in leaf size relative to other plant traits suggest that selection may favour the evolution of larger leaves on islands, perhaps due to release from predation or increased intra‐specific competition.