Diversification shifts in leafroller moths linked to continental colonization and the rise of angiosperms

Tectonic dynamics and niche availability play intertwined roles in determining patterns of diversification. Such drivers explain the current distribution of many clades, whereas events such as the rise of angiosperms can have more specific impacts, such as on the diversification rates of herbivores. The Tortricidae, a diverse group of phytophagous moths, are ideal for testing the effects of these determinants on the diversification of herbivorous clades. To estimate ancestral areas and diversification patterns in Tortricidae, a complete tribal‐level dated tree was inferred using molecular markers (one mitochondrial and five nuclear) and calibrated using fossil constraints. We found that Tortricidae diverged from their sister group c. 120 Myr ago (Ma) and diversified c. 97 Ma, a timeframe synchronous with the rise of angiosperms in the Early–mid Cretaceous. Ancestral areas analysis, based on updated Wallace's biogeographical regions, supports the hypothesis of a Gondwanan origin of Tortricidae in the South American plate. We also detected an increase in speciation rate that coincided with the peak of angiosperm diversification in the Cretaceous. This in turn probably was further heightened by continental colonization of the Palaeotropics when angiosperms became dominant by the end of the Late Cretaceous.     

Biogeographical origins and diversification of the exoneurine allodapine bees of Australia (Hymenoptera,Apidae)

Aim Early diversification of allodapine bees occurred in Africa c. 50 Ma. They are most abundant in sub‐Saharan Africa and Australia, and one of the oldest phylogenetic divergences in the tribe involves a split between an African + Malagasy clade and an Australian clade. The historical biogeographical scenario for this has been highly problematic, entailing an Eocene dispersal from Africa to Australia, followed by an unresolved, and apparently rapid, set of bifurcations leading to the Australian ‘exoneurine’ genera. Here we use an expanded taxon set of Australian species to explore the timing and historical biogeography of the exoneurine radiation. Location Australia, Africa, Madagascar. Methods One nuclear gene (F2 copy of elongation factor 1α) and two mitochondrial genes (cytochrome c oxidase subunit I and cytochrome b) were sequenced for 33 Australian exoneurine species from all five genera found on the continent, as well as for an additional 37 species from all non‐parasitic genera in the remainder of the tribe. We used Bayesian inference analyses to study phylogenetic topology and penalized likelihood analyses to infer key dates of divergence within the tribe. We also used lineage‐through‐time (LTT) analyses and Bayesian analyses to explore the tempo of radiations and biogeographical history of the exoneurines. Results Results from the phylogenetic analyses were congruent with previous studies, indicating a single colonization event c. 34 Ma, too late for Gondwanan vicariance models, and too early for a Laurasian dispersal route. In contrast to earlier studies, we show that this colonization event did not result in an ancient rapid radiation. However, LTT patterns indicated a rapid radiation of the temperate‐adapted genera Exoneura and Brevineura, but not of the xeric‐adapted genus Exoneurella, from 10 to 6 Ma. Main conclusions Our results indicate a trans‐oceanic dispersal event from Africa to Australia, most likely via Antarctica, with an accelerated diversification of temperate‐adapted lineages during the major Late Miocene event referred to as the ‘Hill Gap’. This is the first study to link radiations in Australian bee faunal elements to changing climate, and differs from many other plant and insect phylogenetic studies by showing increased radiation of temperate clades, rather than xeric clades, with increasing aridification of Australia.     

100 million years of morphological conservation in bark beetles (Coleoptera: Curculionidae: Scolytinae)

Abstract Scolytine weevils (bark and ambrosia beetles) have a unique ecological significance in forest ecosystems, which equates to major effects on landscape ecology and to monetary losses. Fossilized galleries of scolytines have been reported in Late Mesozoic wood, but here we describe a well‐preserved body fossil from the Cretaceous, c. 100 Ma, preserved in amber from northern Myanmar. Moreover, the specimen is remarkably similar to Recent species of the genus Microborus, revealing stasis unexpected within scolytines and thus highlighting the antiquity of the group. Stratigraphic dating and comparison of insect palaeofaunas included in other well‐dated ambers from multiple sites support the age estimate of the Burmese amber. A minimum age for one clade of scolytines is thus established, indicating an early divergence of scolytines from other weevils in the Late Jurassic or Early Cretaceous and challenging the current perspective of weevil evolution.     

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.     

Population structure within an alpine archipelago: strong signature of past climate change in the New Zealand rock wren (Xenicus gilviventris)

Naturally subdivided populations such as those occupying high‐altitude habitat patches of the ‘alpine archipelago’ can provide significant insight into past biogeographical change and serve as useful models for predicting future responses to anthropogenic climate change. Among New Zealand's alpine taxa, phylogenetic studies support two major radiations: the first correlating with geological forces (Pliocene uplift) and the second with climatic processes (Pleistocene glaciations). The rock wren (Xenicus gilviventris) is a threatened alpine passerine belonging to the endemic New Zealand wren family (Acanthisittidae). Rock wren constitute a widespread, naturally fragmented population, occurring in patches of suitable habitat over c. 900 m in altitude throughout the length of the South Island, New Zealand. We investigated the relative role of historical geological versus climatic processes in shaping the genetic structure of rock wren (N = 134) throughout their range. Using microsatellites combined with nuclear and mtDNA sequence data, we identify a deep north–south divergence in rock wren (3.7 ± 0.5% at cytochrome b) consistent with the glacial refugia hypothesis whereby populations were restricted in isolated refugia during the Pleistocene c. 2 Ma. This is the first study of an alpine vertebrate to test and provide strong evidence for the glacial refugia hypothesis as an explanation for the low endemicity central zone known as the biotic ‘gap’ in the South Island of New Zealand.     

A revision of the sauropod dinosaur genus ‘Bothriospondylus’ with a redescription of the type material of the Middle Jurassic form ‘B. madagascariensis’

Abstract: The sauropod dinosaur ‘Bothriospondylus’, originally named on the basis of Late Jurassic remains from England, is demonstrated to be invalid, and the characters used to diagnose it are shown to be obsolescent features which are widespread throughout Sauropoda. Material referred to this genus spans a temporal range from the Middle Jurassic until the early Late Cretaceous and has been described from five different countries, across three continents. These remains represent a wide array of sauropod groups, comprising non‐neosauropod eusauropods, a macronarian, titanosauriforms (including at least one definite brachiosaurid) and a rebbachisaurid. The type material of the Middle Jurassic ‘B. madagascariensis’ represents a derived non‐neosauropod eusauropod and possesses two potential autapomorphies. However, as a result of the fragmentary nature of the material and the uncertainty surrounding its association, a new taxon is not erected. Of the numerous specimens referred to ‘Bothriospondylus’, however, several remains are considered diagnostic: Ornithopsis hulkei (Early Cretaceous, UK), Lapparentosaurus madagascariensis (Middle Jurassic, Madagascar) and Nopcsaspondylus alarconensis (early Late Cretaceous, Argentina). At least three types of sauropod were present in the Bathonian (Middle Jurassic) of north‐west Madagascar, with a basal eusauropod (Archaeodontosaurus), a more derived eusauropod (‘B. madagascariensis’) and a titanosauriform (Lapparentosaurus) all approximately contemporaneous. Palaeocontinental reconstructions suggest that Middle Jurassic Madagascan sauropods would still have been capable of global biotic interchange, and this is perhaps reflected in their diverse assemblage. Re‐evaluation of these Malagasy forms has shed new light on this important time period in sauropod evolution.     

The oldest higher true crabs (Crustacea: Decapoda: Brachyura): insights from the Early Cretaceous of the Americas

Despite the extensive fossil record of higher crabs (Eubrachyura) from Late Cretaceous and Cenozoic rocks worldwide, their Early Cretaceous occurrences are scarce and fragmentary, obscuring our understanding of their early evolution. Until now, representatives of only two families of eubrachyuran‐like crabs were known from the Early Cretaceous: Componocancridae and Tepexicarcinidae fam. nov., both monospecific lineages from the Albian (~110–100 Ma) of North and Central America, respectively. The discovery of Telamonocarcinus antiquus sp. nov. (Telamonocarcinidae) from the early Albian of Colombia, South America (~110 Ma), increases to three the number of known Early Cretaceous eubrachyuran‐like families. The ages and geographical distributions of the oldest eubrachyuran‐like taxa (i.e. Componocancridae, Telamonocarcinidae and Tepexicarcinidae fam. nov.) suggest that the oldest higher true crabs might have originated in the Americas; that they were already morphologically diverse by the late Early Cretaceous; and that their most recent common ancestor must be rooted in the Early Cretaceous, or even the Late Jurassic.     

Gondwanan radiation of the Southern Hemisphere crayfishes (Decapoda: Parastacidae): evidence from fossils and molecules

Aim The sequential break‐up of Gondwana is thought to be a dominant process in the establishment of shared biota across landmasses of the Southern Hemisphere. Yet similar distributions are shared by taxa whose radiations clearly post‐date the Gondwanan break‐up. Thus, determining the contribution of vicariance versus dispersal to seemingly Gondwanan biota is complex. The southern freshwater crayfishes (family Parastacidae) are distributed on Australia and New Guinea, South America, Madagascar and New Zealand and are unlikely to have dispersed via oceans, owing to strict freshwater limitations. We test the hypotheses that the break‐up of Gondwana has led to (1) a predominately east–west (((Australia, New Zealand: 80 Ma) Madagascar: 160–121 Ma) South America: 165–140 Ma), or (2) a southern (((Australia, South America: 52–35 Ma) New Zealand: 80 Ma) Madagascar: 160–121 Ma) pattern for parastacid crayfish. Further, we examine the evidence for a complete drowning of New Zealand and subsequent colonization by freshwater crayfish. Location Southern Hemisphere. Methods The evolutionary relationships among the 15 genera of Parastacidae were reconstructed using mitochondrial [16S, cytochrome c oxidase subunit I (COI)] and nuclear (18S, 28S) sequence data and maximum likelihood and Bayesian methods of phylogenetic reconstruction. A Bayesian (multidivtime ) molecular dating method using six fossil calibrations and phylogenetic inference was used to estimate divergence time among crayfish clades on Gondwanan landmasses. Results The South American crayfish are monophyletic and a sister group to all other southern crayfish. Australian crayfish are not monophyletic, with two Tasmanian genera, Spinastacoides and Ombrastacoides, forming a clade with New Zealand and Malagasy crayfish (both monophyletic). Divergence of crayfish among southern landmasses is estimated to have occurred around the Late Jurassic to Early Cretaceous (109–178 Ma). Main conclusions The estimated phylogenetic relationships and time of divergence among the Southern Hemisphere crayfishes were consistent with an east–west pattern of Gondwanan divergence. The divergence between Australia and New Zealand (109–160 Ma) pre‐dated the rifting at around 80 Ma, suggesting that these lineages were established prior to the break‐up. Owing to the age of the New Zealand crayfish, we reject the hypothesis that there was a complete drowning of New Zealand crayfish habitat.     

Biogeographical and geological evidence for a smaller, completely-enclosed Pacific Basin in the Late Cretaceous

Aim To use biogeographical, palaeomagnetic, palaeosedimentary, and plate circuit data from Late Cretaceous regions in and around the Pacific to test the plate tectonic hypothesis of a pre‐Pacific superocean. Location East Asia, Australia, Antarctica, the western Americas, and the Pacific. Methods Literature surveys of the distributions of Cretaceous, circum‐Pacific taxa were compared with palaeomagnetic and palaeosedimentary data. Uncontroversial plate motions based on seafloor spreading data were also used to test the results of the biogeographical and palaeomagnetic analyses. Results The distributions of Cretaceous terrestrial taxa, mostly dinosaurs, imply direct, continental connections between Australia and East Asia, East Asia and North America, North America and South America, South America and Antarctica, and Antarctica and Australia. Palaeomagnetic, palaeosedimentary, and basic plate circuit analyses require little to no latitudinal motion of the Pacific plate with respect to the surrounding continents. Specifically, the data implies that western North America, East Asia, and the Pacific plate all increased in latitude by roughly the same amount (c. 11 ± 5°) since the Campanian – and that the Pacific Ocean Basin has increased in length north‐to‐south. Main conclusions Each of the analyses provides independent corroboration for the same conclusion: the Late Cretaceous Pacific plate was completely enclosed by the surrounding continents and has not experienced significant latitudinal motion with respect to North America, East Asia, or the Bering land bridge. This contrasts significantly with the plate tectonic history of the Pacific, implying instead that the Pacific plate formed in situ, pushing the continents apart as the plate and basin expanded. These results also substantiate recent biogeographical analyses that have concluded that a narrower Pacific Ocean Basin in the Mesozoic and early Tertiary provides the most reasonable explanation for the great number of trans‐Pacific disjunctions of poor dispersing taxa.     

Lygosomine phylogeny and the origins of Australian scincid lizards

Aim Australian scincid lizards represent three distinct groups within the cosmopolitan clade Lygosominae, the Egernia, Eugongylus and Sphenomorphus groups. This paper presents a time‐calibrated phylogeny for Lygosominae that provides the necessary temporal framework for assessing the contributions of immigration from Asia and of Gondwanan inheritance in the derivation of the Australian scincid fauna. Location Australasia, Asia, Africa. Methods Phylogenetic relationships and divergence times were inferred from novel BDNF, c‐mos and PTPN12 sequences (2408 aligned sites). Results Lygosomine monophyly is well supported, and there is strong support for monophyly of the Egernia, Eugongylus and Sphenomorphus groups. A sister‐group relationship of Tribolonotus (distributed in Melanesia and the Papuan Region) and the Egernia group is strongly supported in both Bayesian and maximum likelihood analyses. Australian representatives of the Sphenomorphus group compose a significantly supported clade estimated to have originated c. 25 Ma. An age of c. 18 Ma is inferred for a strongly supported clade comprising Australian representatives of the Egernia group; this clade diverged from Corucia zebrata (confined to the Solomon Islands) c. 25 Ma and from Tribolonotus c. 54 Ma. A well‐supported clade including all Australian Eugongylus group taxa sampled is estimated to have arisen c. 20 Ma. Main conclusions The Australian Sphenomorphus group is nested within the more inclusive Sphenomorphus group (distributed primarily in Asia and Australasia), suggesting comparatively recent descent from a colonizing Asian ancestor; the divergence times inferred here indicate that colonization occurred during the mid Cenozoic, subsequent to the rifting of Australia from Antarctica. An Oligocene origin of the extant Eugongylus group fauna of Australasia (the basal members of which are distributed in the Southwest Pacific) indicates that Eugongylus group lygosomines also dispersed to Australia relatively recently. The Egernia group diverged from Tribolonotus in the Early Eocene; however, extant Egernia group lygosomines originated only during the Late Oligocene, implying extensive pruning of stem taxa (i.e. extinction). As a result, inferences of the timing of dispersal into Australia are associated with substantial uncertainty, although independent palaeontological evidence suggests that the Egernia group entered Australia prior to the Oligocene, immediately after (or perhaps before) the separation of Australia and Antarctica.     

A Plains‐wanderer (Pedionomidae) that did not wander plains: a new species from the Oligocene of South Australia

The remarkable fauna of Australia evolved in isolation from other landmasses for millions of years, yet understanding the evolutionary history of endemic avian lineages on the continent is confounded by the ability of birds to disperse over geographical barriers even after vicariance events. The Plains‐wanderer Pedionomus torquatus (Charadriiformes) is an enigmatic, predominantly sedentary, quail‐like bird that occurs exclusively in sparse native grasslands of southeastern Australia. It is the only known species of its family (Pedionomidae), and its closest relatives are the South American seedsnipes (Thinocoridae). Here we describe a further representative of this lineage, Oligonomus milleri gen. et sp. nov., from the Late Oligocene of South Australia (26–24 Ma), which pre‐dates the earliest record of P. torquatus by c. 22 Ma and attests to the presence of this lineage during Australia's period of isolation (50–15 Ma). Based on the morphology of the coracoid and the palynological record, we propose that O. milleri and P. torquatus were ecologically disparate taxa and that, similar to coeval marsupials, O. milleri inhabited well‐wooded habitats, suggesting that the preference for grassland in the extant P. torquatus and thinocorids is likely to be convergent and not ancestral. The speciation event leading to the evolution of the extant Plains‐wanderer was probably triggered by the spread of grasslands across Australia in the Late Miocene–Pliocene, which this record pre‐dates. The presence of a pedionomid in the Late Oligocene of Australia strengthens the hypothesis of a Gondwanan divergence of the lineages giving rise to Thinocoridae and Pedionomidae.     

Large‐scale molecular phylogeny of Cryptorhynchinae (Coleoptera,Curculionidae) from multiple genes suggests American origin and later Australian radiation

The monophyly of the highly diverse weevil subfamily Cryptorhynchinae is tested with a dataset of 203 taxa representing 159 genera of Curculionoidea, 105 of them Cryptorhynchinae s.l. We construct a phylogeny based on an alignment of 5523 bp, consisting of fragments from two mitochondrial genes (two fragments of COI, 16S) and seven nuclear genes (ArgK, CAD, EF1α, enolase, H4, 18S, 28S). Analyses of maximum likelihood and Bayes inference recovered largely congruent results. Groups with different morphology of the rostral furrow (e.g. Aedemonini, Camptorhinini, Cryptorhynchini, Ithyporini) are not closely related to each other. However, most taxa with a mesosternal receptacle are monophyletic and here defined as Cryptorhynchinae s.s., comprising Cryptorhynchini, Gasterocercini, Torneumatini and Psepholacini, but also Arachnopodini and Idopelma Faust. The genus Phyrdenus LeConte is excluded from Cryptorhynchinae and transferred to Conotrachelini of Molytinae. Thus defined, the group still comprises several thousand species with centres of its diversity in South America and Australia. The early lineages we find in America and the Palearctic, while the extremely diverse faunas of Australia and neighbouring islands mainly belong to a more recent, species‐rich radiation. This also includes a clade comprising the majority of litter‐inhabiting species of New Zealand and the genus Miocalles Pascoe. Flightlessness was attained repeatedly and resulted in convergent evolution of a similar habitus in different zoogeographic regions, mainly exhibited by the polyphyletic genus Acalles Schoenherr.     

Biogeography and diversification of the Pacific ant genus Lordomyrma Emery

Aim This study addresses the origins of terrestrial biodiversity of the Fijian islands using the ant genus Lordomyrma (Hymenoptera: Formicidae: Myrmicinae) as a model system. We derive the evolution of the genus and determine its closest extra-Fijian relatives from geological data, molecular phylogenetic reconstruction and divergence estimates. Location Ant taxa were sampled in the Southwest Pacific, Melanesia, Southeast Asia, Australia and mainland China. Methods Phylogeny and divergence estimates of the ant genus Lordomyrma based on four nuclear genes (28S, ArgK, LW Rh, CAD) plus data on Indo-Pacific geological history are used to address current hypotheses regarding the origins of the Fijian biota. Results The genus Lordomyrma probably originated in mainland Asia, with subsequent colonization of Australia and the Pacific. The Fijian Lordomyrma clade is monophyletic, and originated c. 8.8 Ma, when it diverged from a sister group in Papua New Guinea. Main conclusions The colonization of Fiji by Lordomyrma is probably a result of long-distance dispersal from New Guinea, possibly aided by island hopping across the Vitiaz Arc. The timeline of diversification in Lordomyrma is broadly congruent with the Miocene fragmentation of the Vitiaz Arc and the Pliocene emergence of Vanua Levu. The biotic shuttle hypothesis, which posits ‘Eua Island as the source of Fijian endemics, is rejected based on the sister relationship of Fiji and New Guinea lineages, as well as on the Miocene submergence of the terrane below sea level. The diversity of Fijian Lordomyrma results from the radiation of a single lineage, which diverged from a New Guinea sister group. The genus appears to have originated in Asia rather than in Australia.     

Phylogeography of Sophora davidii (Leguminosae) across the ‘Tanaka‐Kaiyong Line’, an important phytogeographic boundary in Southwest China

The ‘Tanaka‐Kaiyong Line’ (TKL) is a major phytogeographic boundary in Southwest China, separating East Asia's Sino‐Himalayan and Sino‐Japanese Floras. However, little is known about the importance of this boundary in promoting intraspecific phylogeographic subdivision and divergence. Using chloroplast (cpDNA) and nuclear‐intron (nDNA) sequence data, we reconstructed the population history of Sophora davidii, a drought‐tolerant riparian shrub widely distributed on either side of the TKL. Specifically, we aimed at testing two long‐standing explanations for possible vicariant events across the TKL: (i) Late Pliocene (c. 3 Ma) geological uplift of the eastern Qinghai‐Tibetan Plateau (QTP) or (ii) a sharp environmental gradient associated with the establishment of different monsoon regimes on either side of the TKL during the (Late) Pleistocene. Our genealogical analyses detected a major west–east split in cpDNA, geographically largely consistent with the TKL, and dated to c. 1.28 Ma (95% HPD: 0.21–2.96 Ma), hence postdating the latest phase of eastern QTP uplift. Furthermore, integrating cpDNA phylogeographic patterns with mismatch analyses, we found multiple refugial isolation and long‐term demographic stability of populations in the west (Hengduan Mountain Range) compared with extensive range expansions in the east, possibly during the last glacial period(s) and followed by differentiation into regional sublineages (southeast: Yunnan‐Guizhou Plateau vs. northeast: Qinling Mts./Loess Plateau). Although nuclear differentiation was less marked, the geographical pattern of nDNA haplotypes provided some further indication of the species' eastward expansion, possibly from source populations located just east of the TKL (lower Jinshajiang region). Overall, the present data reject the geological (tectonic) explanation for the TKL and, instead, provide supportive evidence for its role as a climatically driven barrier to present‐day plant dispersal. In addition, our study highlights changing temperatures and vegetation types during the last glacial period(s), along with aspects of regional topography, to be important determinants of the glacial eastward expansion of S. davidii. In consequence, our study lends support to a ‘glacial out‐of‐Hengduan Mts’. hypothesis for the xerophytic‐riparian flora of Southwest China, which in turn is inconsistent with the traditional view of the TKL as a ‘classical’ vicariant‐biogeographic boundary.     

Out of the Neotropics: Late Cretaceous colonization of Australasia by American arthropods

The origins of tropical southwest Pacific diversity are traditionally attributed to southeast Asia or Australia. Oceanic and fragment islands are typically colonized by lineages from adjacent continental margins, resulting in attrition of diversity with distance from the mainland. Here, we show that an exceptional tropical family of harvestmen with a trans-Pacific disjunct distribution has its origin in the Neotropics. We found in a multi-locus phylogenetic analysis that the opilionid family Zalmoxidae, which is distributed in tropical forests on both sides of the Pacific, is a monophyletic entity with basal lineages endemic to Amazonia and Mesoamerica. Indo-Pacific Zalmoxidae constitute a nested clade, indicating a single colonization event. Lineages endemic to putative source regions, including Australia and New Guinea, constitute derived groups. Divergence time estimates and probabilistic ancestral area reconstructions support a Neotropical origin of the group, and a Late Cretaceous (ca 82 Ma) colonization of Australasia out of the Fiji Islands and/or Borneo, which are consistent with a transoceanic dispersal event. Our results suggest that the endemic diversity within traditionally defined zoogeographic boundaries might have more complex evolutionary origins than previously envisioned.     

Global historical biogeography of hadrosaurid dinosaurs

Hadrosaurids were the most derived ornithopods and amongst the most diverse herbivore dinosaurs during the Late Cretaceous of Europe, Asia, and the two Americas. Here, their biogeographical history is reconstructed using dispersal‐vicariance analysis (DIVA). The results showed that Hadrosauridae originated in North America and soon after dispersed to Asia no later than the Late Santonian. The most recent common ancestor of Saurolophidae (= Saurolophinae + Lambeosaurinae) is inferred to have been widespread in North America and Asia. The split between saurolophines and lambeosaurines occurred in response to vicariance no later than the Late Santonian: the former clade originated in North America, whereas the latter did so in Asia. Saurolophine biogeographical history included a minimum of five dispersal events followed by vicariance. Four of these dispersals were inferred to have occurred from North America to Asia during the Campanian and Early Maastrichtian, whereas a fifth event represented a southward dispersal from North to South America no later than the Late Campanian. The historical biogeography of lambeosaurines was characterized by an early evolution in Asia, with a Campanian dispersal to the European archipelago followed by vicariance. Reconstruction of the ancestral areas for the deepest nodes uniting the more derived lambeosaurines clades (‘hypacrosaurs’, ‘corythosaurs’, and ‘parasaurolophs’) is ambiguous. The split between North American and Asian clades of ‘hypacrosaurs’ and ‘parasaurolophs’ occurred in response to vicariance during the Campanian. The evolutionary history of North American ‘hypacrosaurs’ and ‘parasaurolophs’ was characterized by duplication events. The latter also characterized the Late Campanian ‘corythosaurs’, which remained restricted to North America. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 503–525.     

Modern hydrophilid clades present and widespread in the Late Jurassic and Early Cretaceous (Coleoptera: Hydrophiloidea: Hydrophilidae)

We present a summary of the fossil evidence documenting the worldwide occurrence of the family Hydrophilidae (Insecta: Coleoptera: Polyphaga: Hydrophiloidea) in the Late Jurassic and Early Cretaceous. We present the first known fossils of the family from the Mesozoic, being c. 100 Myr older than the fossil record available until now. Two Late Jurassic fossils are documented: P rotochares brevipalpis gen. nov., sp. nov. from the Talbragar Fish Bed (New South Wales, Australia) and ‘Mesosperchus’ schultzi Ponomarenko, 1985 from Solnhofen (Bavaria, Germany). The occurrence of the Hydrophilidae in the Early Cretaceous is documented by six species, all of which may be already assigned to modern subfamilies/tribes: B aissalarva hydrobioides gen. nov., sp. nov. from the Baissa outcrops (Buryat Republic, Russia) and C retoxenus australis gen. nov., sp. nov. from Koonwarra outcrops (Victoria, Australia) are both assigned to the tribe Hydrobiusini (Hydrophilinae); A legorius yixianus gen. nov., sp. nov. and Alegorius sp. from the Yixian Formation (Liaoning, China) may represent the Acidocerinae or Enochrinae, H ydroyixia elongata gen. nov., sp. nov. and H . latissima sp. nov. from the same locality are assigned to the Acidocerinae. The phylogenetic position of Baissalarva hydrobioides is also tested by a phylogenetic analysis. The presence of extant clades (Hydrophilinae: Hydrobiusini, Acidocerinae) in the Early Cretaceous and the wide distribution of the Hydrobiusini in both Gondwana and Laurasia at the same time suggests that the principal extant clades of the Hydrophilidae are at least of Early–Middle Jurassic origin. © 2014 The Linnean Society of London     

Cretaceous opine bivalves from the Pacific slope of North America and palaeobiogeography of subfamily Opinae Chavan, 1969

Abstract: Opines constitute a small subfamily of Mesozoic astartid bivalves that lived mostly in the Tethys Sea region. They first appeared in western Europe during the Middle Triassic, became most widespread during the Jurassic and had their first undoubted appearance in the New World during the Late Jurassic. Their Pacific slope of North America record is studied in detail for the first time. The earliest of the Pacific slope opines is Opis californica Stanton of Early Cretaceous (Hauterivian or Barremian) age. The other six opine species in the study area are of Late Cretaceous age (collectively Middle Turonian – early Late Maastrichtian), have hinges bearing two strong cardinal teeth in each valve, and are placed in the new subgenus Hesperopsis, which is the first astartid to have a partially internal ligament. These six species comprise two morphological lineages inferred to have evolved from O. (H.) popenoei sp. nov. The ‘holzana’ lineage includes O. (H.) holzana sp. nov. and O. (H.) rosarioensis Anderson and Hanna. The ‘anae’ lineage includes O. (H.) anae sp. nov., O. (H.) vancouverensis Whiteaves and O. (H.) triangulata (Cooper). Hesperopis lived in warm temperate waters in offshore shelfal areas. It had a vertical commissure and probably lived as an edgewise recliner. Through time, the species of both lineages became larger and either broader or more elongate. Neither Opis shastalis Anderson, of Aptian age, nor Opis virginalis Waring, of late Palaeocene age, are opines. A junior secondary homonym of Opis trigonalis (Sowerby) was detected and is renamed here as Opis tamurai nom nov.     

Out of Australia and back again: the world‐wide historical biogeography of non‐pollinating fig wasps (Hymenoptera: Sycophaginae)

Aim Figs (Ficus, Moraceae) are exploited by rich communities of often host‐specific phytophagous wasps. Among them, gall‐inducing Sycophaginae (Hymenoptera, Chalcidoidea) may share a common history with Ficus and their mutualistic pollinators (Agaonidae). We investigate here, for the first time, the phylogeny and biogeographical history of Sycophaginae and compare the timing of radiation and dispersion of major clades with available data on Ficus and fig pollinators. Reconstructing the history of their host colonization and association over space and time is central to understanding how fig wasp communities were assembled. Location World‐wide. Methods Maximum likelihood and Bayesian analyses were conducted on 4267 bp of mitochondrial and nuclear DNA to produce a phylogeny of all genera of Sycophaginae. Two relaxed clock methods with or without rate autocorrelation were used for date estimation. Analyses of ancestral area were also conducted to investigate the geographical origin of the Sycophaginae. Results The phylogeny is well resolved and supported. Our data suggest a post‐Gondwanan origin for the Sycophaginae (50–40 Ma) and two independent out‐of‐Australia dispersal events to continental Asia. Given palaeoclimatic and palaeogeographic records, the following scenario appears the most likely. The ancestor of Idarnes+Apocryptophagus migrated to Greater India through the Ninetyeast Ridge (40–30 Ma). The ancestor of Anidarnes+Conidarnes dispersed later via Sundaland (25–20 Ma). Idarnes and Anidarnes subsequently reached the New World via the North Atlantic land bridges during the Late Oligocene Warming Event. Apocryptophagus reached Africa c. 20 Ma via the Arabic corridors and returned to Australasia following the expansion of Sundaland tropical forests (20–10 Ma). Main conclusions Sycophaginae probably invaded the fig microcosm in Australia c. 50–40 Ma after the origin of their host plant. Once associated with figs, they dispersed out of Australia and radiated together with their host fig and associated pollinator through the tropics. We recorded a good coincidence of timing between dispersal events of Sycophaginae and continental connections. Furthermore, fruit pigeons that disperse figs probably spread out of Australasia through the Indian Ocean via the Ninetyeast Ridge c. 38 Ma. Therefore, our study highlights the potential for combining molecular phylogenetics with multiple methods of dating of interacting groups to reconstruct the historical biogeography of plant–herbivore associations.     

Pronemouridae fam. nov. (Insecta: Plecoptera), the stem group of nemouridae and notonemouridae,from the Middle Jurassic of Inner Mongolia,China

Abstract: A new family, Pronemouridae fam. nov., with a new genus, Pronemoura gen. nov., and five new species of fossil stoneflies are described from Daohugou village (Middle Jurassic), Inner Mongolia, China: P. shii sp. nov., P. angustithorax sp. nov., P. longialata sp. nov., P. minuta sp. nov. and P. peculiaris sp. nov. Wing venation of pronemourinids presents some or considerable variety in the fore and hind wings of the same or different individuals. Pronemourinidae, retaining plesiomorphic characters (short, multisegmented cerci and CuA with forks), should be treated as the stem group of Nemouridae and Notonemouridae. Based on fossil data, we propose a model in which the extant plecopterid distribution began in the Early Cretaceous. Nemouridae and Notonemouridae must have occurred in Asia and begun to migrate from here by at least the Early Cretaceous.