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.     

The timing of diversification within the most divergent parrot clade

The Strigopidae are an ancient parrot (Psittaciformes) family consisting of three extant species placed in two genera (Nestor, Strigops) and restricted to New Zealand. Their evolutionary history is clouded because the timing of divergence events within this family has variously been attributed to Pleistocene climate change or much earlier earth‐historic events. Here we examine new psittaciform DNA sequence data, and combine them with previously published sequences, to shed light on the poorly understood timing of diversification within the Strigopidae. Using calibrations indirectly derived from both psittaciform and non‐psittaciform fossils, our data indicate a Late Pliocene or Early Pleistocene (ca 1.2–3.6 mya) differentiation between the two Nestor species (kea and kaka), possibly in response to shifts in habitat distribution associated with sea level fluctuations. The unique, monotypic, nocturnal and flightless genus Strigops (kakapo) is shown to have diverged from the Nestor lineage probably ca 28–29 mya, coinciding with the potential Oligocene submergence of Zealandia when much of its landmass may have been fragmented into smaller islands, providing a setting for allopatric diversification.     

Support for vicariant origins of the New Zealand Onychophora

Aim The distribution of Onychophora across the southern continents has long been considered the result of vicariance events. However, it has recently been hypothesized that New Zealand was completely inundated during the late Oligocene (25–22 Ma) and therefore that the entire biota is the result of long-distance dispersal. We tested this assumption using phylogenetic and molecular dating of DNA sequence data from Onychophora. Location New Zealand, Australia, South Africa, Chile (South America). Methods We obtained DNA sequence data from the nuclear genes 28S and 18S rRNA to reconstruct relationships among species of Peripatopsidae (Onychophora). We performed molecular dating under a Bayesian relaxed clock model with a range of prior distributions using the rifting of South America and South Africa as a calibration. Results Our phylogenetic trees revealed that the New Zealand genera Ooperipatellus and Peripatoides, together with selected Australian genera (Euperipatoides, Phallocephale and an undescribed genus from Tasmania), form a monophyletic group that is the sister group to genera from Chile (Metaperipatus) and South Africa (Peripatopsis and Opisthopatus). The relaxed clock dating analyses yielded mean divergence times from 71.3 to 78.9 Ma for the split of the New Zealand Peripatoides from their Australian sister taxa. The 0.95 Bayesian posterior intervals were very broad and ranged from 24.5 to 137.6 Ma depending on the prior assumptions. The mean divergence of the New Zealand species of Ooperipatellus from the Australian species Ooperipatellus insignis was estimated at between 39.9 and 46.2 Ma, with posterior intervals ranging from 9.5 to 91.6 Ma. Main conclusions The age of Peripatoides is consistent with long-term survival in New Zealand and implies that New Zealand was not completely submerged during the Oligocene. Ooperipatellus is less informative on the question of continuous land in the New Zealand region because we cannot exclude a post-Oligocene divergence. The great age of Peripatoides is consistent with a vicariant origin of this genus resulting from the rifting of New Zealand from the eastern margin of Gondwana and supports the assumptions of previous authors who considered the Onychophora to be a relict component of the New Zealand biota.     

A new species of watering pot shell (Bivalvia: Anomalodesmata: Clavagelloidea) from the Miocene of the Murray Basin,South Australia

Abstract: A new species of penicillid watering pot shell, Kendrickiana coquinacola sp. nov., is described from the middle Miocene (Balcombian) Bryant Creek Formation of the Murray Basin, South Australia. The new species differs from the extant K. veitchi in its smaller size, much shorter posterior tube, fewer tubules in the anterior watering pot structure, absence of the pedal slit, discontinuous dorsolateral bands of pitted muscle scars on the internal surface of the anterior bulb and habit of cementing itself to the shells in its surrounding environment. The fossil record of Kendrickiana is reviewed. The record from the Dry Creek Sands is discounted, while a record for the extant K. veitchi from the earliest Pleistocene of York Peninsula is added. K. coquinacola indicates the highly derived anatomy of the genus evolved over a 10‐Ma period from the late Oligocene through the early Miocene.     

After the deluge: mitochondrial DNA indicates Miocene radiation and Pliocene adaptation of tree and giant weta (Orthoptera: Anostostomatidae)

Aim New Zealand broke away from the margins of Gondwana c. 75 Ma. Since then, New Zealand taxa derived from the Gondwanan biota are thought to have been exposed first to a subtropical climate on a low lying terrain, then severe land reduction during the Oligocene marine transgression, followed by much cooler climates of the Pliocene and Pleistocene, at which time mountain ranges emerged. The biological consequence of New Zealand's geological and climatic history is not well understood, in particular the extent to which the Oligocene acted as a biological bottleneck remains unresolved. Methods We used mitochondrial cytochrome oxidase I and 12S DNA sequences to examine the extent of diversity and inferred timing of speciation of New Zealand weta (Anostostomatidae), a group of Orthoptera with a Gondwanan distribution generally thought to be ancient inhabitants of New Zealand. Main conclusions We hypothesize that at least three distinct groups of weta survived the Oligocene marine transgression and radiated subsequently. Speciation followed during the Miocene and radiation into new habitats occurred during the Pliocene when mountain building created novel environments. Patterns of genetic diversity within species reflect, in some instances, geographical subdivision in the Pliocene, and in other cases, Pleistocene range changes resulting from climate change.     

Afrotropical and Nearctic genera of Odonata in the French Oligocene: biogeographic and paleoclimatic implications (Insecta: Calopterygidae,Aeshnidae)

New species of the genera Sapho and Epiaeschna are recorded in the Oligocene of Aix- en-Provence, Bouches-du-Rhône, France. Sapho legrandi n. sp. is the third fossil representative of this recent African genus and Epiaeschna pseudoheros n. sp. is the fifth fossil species of this recent North American genus. The fossil species Triaeschna gossi from the Eocene of England, Epacantha magnifica from the Late Oligocene of Kazakhstan, and Mediaeschna matutina from the Oligocene of China, are considered species of Epiaeschna and the three fossil genera Triaeschna Campion 1916, Mediaeschna Zhang 1989, and Epacantha Martynov 1929 are synonymized with Epiaeschna. The closely related genera Umma and Sapho inhabit warm humid forests of Western Africa. Their presence in two Oligocene deposits of France supports the hypothesis of a warm humid palaeoenvironment for Armissan (Aude, France), and Aix-en-Provence.     

Out of Africa: Biogeography and diversification of the pantropical pond skater genus Limnogonus Stål, 1868 (Hemiptera: Gerridae)

Gondwanan vicariance, long‐distance dispersal (LDD), and boreotropical migration have been proposed as alternative hypotheses explaining the pantropical distribution pattern of organisms. In this study, the historical biogeography of the pond skater genus Limnogonus was reconstructed to evaluate the impact of biogeographical scenarios in shaping their modern transoceanic disjunction. We sampled almost 65% of recognized Limnogonus species. Four DNA fragments including 69 sequences were used to reconstruct a phylogram. Divergence time was estimated using a Bayesian relaxed clock method and three fossil calibrations. Diversification dynamics and ancestral area reconstruction were investigated by using maximum likelihood and Bayesian approaches. Our results showed the crown group of Limnogonus originated and diversified in Africa in the early Eocene (49 Ma, HPD: 38–60 Ma), subsequently expanding into other regions via dispersal. The colonization of the New World originated from the Oriental Region probably via the Bering Land Bridge in the late Eocene. Two split events between the Old World and New World were identified: one between Neotropics and Oriental region around the middle Oligocene (30 Ma, HPD: 22–38 Ma), and the other between Neotropics and Africa during the middle Miocene (14 Ma, HPD: 8–21 Ma). The evolutionary history of Limnogonus involved two biogeographical processes. Gondwanan vicariance was not supported in our analyses. The diversification of Limnogonus among Africa, Oriental, and Neotropical regions corresponded with the age of land bridge connection and dispersed as a member associated with the broad boreotropical belt before local cooling (34 Ma). The current transoceanic disjunctions in Limnogonus could be better explained by the disruption of “mixed‐mesophytic” forest belt; however, the direct transoceanic LDD between the Neotropics and Africa could not be ruled out. In addition, the “LDD” model coupled with island hopping could be a reasonable explanation for the diversification of the Oriental and Australian regions during the Oligocene.     

Radiolarian faunal turnover across the Oligocene/Miocene boundary in the equatorial Pacific Ocean

The global warming trend of the latest Oligocene was interrupted by several cooling events associated with Antarctic glaciations. These cooling events affected surface water productivity and plankton assemblages. Well-preserved radiolarians were obtained from upper Oligocene to lower Miocene sediments at Ocean Drilling Program (ODP) Leg 199 Sites 1218 and 1219 in the equatorial Pacific, and 110 radiolarian species were identified.Four episodes of significant radiolarian faunal changes were identified: middle late Oligocene (27.5 to 27.3 Ma), latest Oligocene (24.4 Ma), earliest Miocene (23.3 Ma), and middle early Miocene (21.6 Ma). These four episodes approximately coincide with increases and decreases of biogenic silica accumulation rates and increases in δ¹⁸O values coded as “Oi” and “Mi” events. These data indicate that Antarctic glaciations were associated with change of siliceous sedimentation patterns and faunal changes in the equatorial Pacific.Radiolarian fauna was divided into three assemblages based on variations in radiolarian productivity, species richness and the composition of dominant species: a late Oligocene assemblage (27.6 to 24.4 Ma), a transitional assemblage (24.4 to 23.3 Ma) and an early Miocene assemblage (23.3 to 21.2 Ma). The late Oligocene assemblage is characterized by relatively high productivity, low species richness and four dominant species of Tholospyris anthophora, Stichocorys subligata, Lophocyrtis nomas and Lithelius spp. The transitional assemblage represents relatively low values of productivity and species richness, and consists of three dominant species of T. anthophora, S. subligata and L. nomas. The characteristics of the early Miocene assemblage are relatively low productivity, but high species richness. The two dominant species present in this assemblage are T. anthophora and Cyrtocapsella tetrapera. The most significant faunal turnover of radiolarians is marked at the boundary between the transitional/early Miocene assemblages.We also reviewed changes in other microfossil assemblages in the low latitudes during the late Oligocene through early Miocene. The microfossil assemblages of major groups show sequential changes near the Oligocene/Miocene (O/M) boundary (23.8 Ma). Many extinction events and some first occurrences of calcareous nannofossils and many occurrences of radiolarians are found from about 24.8 to 23.3 Ma, and first occurrences of planktic foraminifers and diatoms followed from 23.2 through 22 Ma. Hence, the O/M boundary is identified as a significant level for microfossil evolutions.     

Small mammal (rodents and lagomorphs) European biogeography from the Late Oligocene to the mid Pliocene

Aim To analyse the fossil species assemblages of rodents and lagomorphs from the European Neogene in order to assess what factors control small mammal biogeography at a deep‐time evolutionary time‐scale. Location Western Europe: 626 fossil‐bearing localities located within 31 regions and distributed among 18 successive biochronological units ranging from c. 27 Ma (million years ago; Late Oligocene) to c. 3 Ma (mid Pliocene). Methods Taxonomically homogenized pooled regional assemblages are compared using the Raup and Crick index of faunal similarity; then, the inferred similarity matrices are visualized as neighbour‐joining trees and by projecting the statistically significant interregional similarities and dissimilarities onto palaeogeographical maps. The inferred biogeographical patterns are analysed and discussed in the light of known palaeogeographical and palaeoclimatic events. Results Successive time intervals with distinct biogeographical contexts are identified. Prior to c. 18 Ma (Late Oligocene and Early Miocene), a relative faunal homogeneity (high interregional connectivity) is observed all over Europe, a time when major geographical barriers and a weak climatic gradient are known. Then, from the beginning of the Middle Miocene onwards, the biogeography is marked by a significant decrease in interregional faunal affinities which matches a drastic global climatic degradation and leads, in the Late Miocene (c. 11 Ma), to a marked latitudinal pattern of small mammal distribution. In spite of a short rehomogenization around the Miocene/Pliocene boundary (6–4 Ma), the biogeography of small mammals in the mid Pliocene (c. 3 Ma) finally closely reflects the extant situation. Main conclusions The resulting biogeographical evolutionary scheme indicates that the extant endemic situation has deep historical roots corresponding to global tectonic and climatic events acting as primary drivers of long‐term changes. The correlation of biogeographical events with climatic changes emphasizes the prevalent role of the climate over geography in generating heterogeneous biogeographical patterns at the continental scale.     

A bittern (Aves: Ardeidae) from the early Miocene of New Zealand

Herons (Aves: Ardeidae) are rare in the fossil record globally. Fossil taxa referred to Ardeinae and Nycticoracini are known from as early as the early Oligocene and ardeids undetermined to subfamily include some as old as the early Eocene. In Australasia, the pre-Pliocene record is restricted to one species from the early Miocene of New Zealand. On the basis of a tarsometatarsus and a coracoid we describe a new species of bittern (Ardeidae: Botaurinae) from the St Bathans Fauna, of early Miocene age, from Otago, New Zealand. This is only the third and the oldest pre-Quaternary record for Botaurinae globally.     

Evolution of New Zealand's terrestrial fauna: a review of molecular evidence

New Zealand biogeography has been dominated by the knowledge that its geophysical history is continental in nature. The continental crust (Zealandia) from which New Zealand is formed broke from Gondwanaland ca 80 Ma, and there has existed a pervading view that the native biota is primarily a product of this long isolation. However, molecular studies of terrestrial animals and plants in New Zealand indicate that many taxa arrived since isolation of the land, and that diversification in most groups is relatively recent. This is consistent with evidence for species turnover from the fossil record, taxonomic affinity, tectonic evidence and observations of biological composition and interactions. Extinction, colonization and speciation have yielded a biota in New Zealand which is, in most respects, more like that of an oceanic archipelago than a continent.     

Miocene Fossils Reveal Ancient Roots for New Zealand’s Endemic Mystacina (Chiroptera) and Its Rainforest Habitat

The New Zealand endemic bat family Mystacinidae comprises just two Recent species referred to a single genus, Mystacina. The family was once more diverse and widespread, with an additional six extinct taxa recorded from Australia and New Zealand. Here, a new mystacinid is described from the early Miocene (19–16 Ma) St Bathans Fauna of Central Otago, South Island, New Zealand. It is the first pre-Pleistocene record of the modern genus and it extends the evolutionary history of Mystacina back at least 16 million years. Extant Mystacina species occupy old-growth rainforest and are semi-terrestrial with an exceptionally broad omnivorous diet. The majority of the plants inhabited, pollinated, dispersed or eaten by modern Mystacina were well-established in southern New Zealand in the early Miocene, based on the fossil record from sites at or near where the bat fossils are found. Similarly, many of the arthropod prey of living Mystacina are recorded as fossils in the same area. Although none of the Miocene plant and arthropod species is extant, most are closely related to modern taxa, demonstrating potentially long-standing ecological associations with Mystacina.     

The phylogenetic placement and biogeographical origins of the New Zealand stick insects (Phasmatodea)

The Lanceocercata are a clade of stick insects (Phasmatodea) that have undergone an impressive evolutionary radiation in Australia, New Caledonia, the Mascarene Islands and areas of the Pacific. Previous research showed that this clade also contained at least two of the nine New Zealand stick insect genera. We have constructed a phylogeny of the Lanceocercata using 2277 bp of mitochondrial and nuclear DNA sequence data to determine whether all nine New Zealand genera are indeed Lanceocercata and whether the New Zealand fauna is monophyletic. DNA sequence data were obtained from mitochondrial cytochrome oxidase subunits I and II and the nuclear large subunit ribosomal RNA and histone subunit 3. These data were subjected to Bayesian phylogenetic inference under a partitioned model and maximum parsimony. The resulting trees show that all the New Zealand genera are nested within a large New Caledonian radiation. The New Zealand genera do not form a monophyletic group, with the genus Spinotectarchus Salmon forming an independent lineage from the remaining eight genera. We analysed Lanceocercata apomorphies to confirm the molecular placement of the New Zealand genera and to identify characters that confirm the polyphyly of the fauna. Molecular dating analyses under a relaxed clock coupled with a Bayesian extension to dispersal‐vicariance analysis was used to reconstruct the biogeographical history for the Lanceocercata. These analyses show that Lanceocercata and their sister group, the Stephanacridini, probably diverged from their South American relatives, the Cladomorphinae, as a result of the separation of Australia, Antarctica and South America. The radiation of the New Caledonian and New Zealand clade began 41.06 million years ago (mya, 29.05–55.40 mya), which corresponds to a period of uplift in New Caledonia. The main New Zealand lineage and Spinotectarchus split from their New Caledonian sister groups 33.72 (23.9–45.62 mya) and 29.9 mya (19.79–41.16 mya) and began to radiate during the late Oligocene and early Miocene, probably in response to a reduction in land area and subsequent uplift in the late Oligocene and early Miocene. We discuss briefly shared host plant patterns between New Zealand and New Caledonia. Because Acrophylla sensu Brock & Hasenpusch is polyphyletic, we have removed Vetilia Stål from synonymy with Acrophylla Gray.     

Systematics and evolutionary significance of the small Abrocomidae from the early Miocene of southern South America

Octodontoidea is the most species-rich clade among hystricomorph rodents, and has a fossil record going back to at least the late Oligocene. Affinities of fossils previous to the late Miocene differentiation of the extant families Abrocomidae, Echimyidae and Octodontidae are controversial, essentially because these fossils may share few apomorphies with modern species. In fact, pre-late Miocene representatives of Abrocomidae had not been recognised until very recently. Here we revise the early Miocene genus Acarechimys, originally assigned to Echimyidae, and alternatively to stem Octodontoidea or to Octodontidae. A systematic and parsimony-based phylogenetic analysis of the species traditionally included in Acarechimys showed that this genus is part of stem Abrocomidae. These results are primarily supported by morphology of the mandible and lower molars. Acarechimys is here restricted to three species, A. minutus, A. pulchellus and Acarechimys pascuali sp. nov., while another species, A. constans, is here transferred to a new abrocomid genus. The remaining species were nested within Octodontidae. According to these results, Abrocomidae might have been as diverse as its sister clade Octodontidae-Echimyidae during the late Oligocene–early Miocene. Extinction of this diversity would have resulted in marked loss of evolutionary history, with extant abrocomids being currently restricted to late-diverged euhypsodont representatives.     

Good morning Gondwana

Summary

Vicariance and dispersion both must be considered as possibilities for the fauna and flora of New Zealand and New Caledonia. Oligocene submersion, promoted by the geologists and several biologists, does not seem to have been total. Refuge stations must have existed in mountains and even in plains in some surrounding areas. From there the relicts must have radiated after the partial submersion. Certain “primitive” Chrysomelidae Eumolpinae (Bohumiljania spp.) are closely related to Patagonian genera. Their case is not unique among the terrestrial organisms of New Caledonia. How to explain the occurrence of Amborella in New Caledonia and of the tuataras in New Zealand, already very probably extinct elsewhere during the Paleogene?     

Nothofagus subgenus Brassospora (Nothofagaceae) leaf fossils from New Zealand: a link to Australia and New Guinea?

N othofagus palustris sp. nov. is the first record of well‐preserved leaves of Nothofagus subgenus Brassospora in New Zealand, and is described from an Oligo–Miocene leaf bed in the Gore Lignite Measures of the South Island. Nothofagus palustris is represented by relatively small and variably toothed leaves with cuticular remains that possess all the characteristic features of the subgenus, including the presence of variably sized stomata that are randomly arranged within areoles, hydathodes along the major veins and ‘bulging cells’ within the areoles on the adaxial side. Phylogenetic assessment shows that the leaves are similar to those of Australian Oligocene and Miocene species and may belong to the same clade of Brassospora. Most notably, these species share the derived feature of abundant leaf wax, a feature that is now only well developed in two New Guinean species. This and other evidence allows the possibility that the ancestor of N. palustris reached New Zealand from Australia. However, it is improbable that N. palustris or a similar species was the common ancestor of the clade of Brassospora that is now confined to New Caledonia. Ecologically, N. palustris is unusual among extant and previously described macrofossil species of Brassospora in being found in a relatively open, swampy habitat. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 503–515.