中国东北中生代时期的传粉昆虫与虫媒植物

昆虫与植物的相互作用在昆虫与植物协同进化和发展中起着至关重要的作用。化石记录中发现了昆虫取食花粉、授粉滴、访问中生代裸子植物具胚珠的繁殖器官等多方面的昆虫化石证据。20世纪90年代以来,研究和报道了大量采自中国东北地区保存良好的昆虫与植物化石。例如发表于Science上的2篇具有长喙的双翅目和长翅目昆虫化石文章,暗示着它们很可能参与各种裸子植物的授粉。在本篇文章中,作者回顾了已报道的中国东北晚中生代时期的昆虫化石,并把它们与授粉或访"花"相联系。另外,作者对采自同一层位的同时代的植物化石进行了初步研究并介绍了可能涉及到与昆虫相互作用的代表性植物。今后将进一步深入研究,以期在化石中找到昆虫授粉和访花的直接证据。     

ESTABLISHMENT OF FOSSIL ENTOMOFAUNAS AND THEIR EVOLUTIONARY SUCCESSION IN NORTH CHINA*

Abstract According to a systematic study of large numbers of fossil insects collected from the Paleozoic, Mesozoic and Tertiary strata in North China, three evolutionary mega-stages of fossil insects including 14 fossil entomofaunas and 29 entomassemblages have been established. These insect faunas reflect their order or law of evolution—the history of their rise and fall, and form their evolutionary succession in North China. Such a history provides an important reference not only for understanding fossil insects through time, but also showing for stratigraphical correlation.     

中国北方化石昆虫群的建立及其演化序列(英文)

作者几十年来已积累了众多的中国北方古生代晚期、中生代和第三纪的昆虫化石标本,对其长期的系统研究结果表明：中国北方昆虫化石演化序列可以划分为三个大的演化阶段,即古生代晚期、中生代和第三纪演化阶段,其中包含14个昆虫群虫和29个昆虫组合。这些昆虫群及其组会既反映了北方昆虫在地质历史上兴衰的发展规律,也为地层划分、对比与地质时代的鉴定提供了依据。     

The earliest fossil evidence of bone boring by terrestrial invertebrates,examples from China and South Africa

We report the oldest fossil evidence of osteophagia by terrestrial invertebrates on both the Asian and African continents. Bones attributable to the Middle Jurassic dinosaur Chuanjiesaurus (Dinosauria: Sauropoda) were found with post-mortem insect modification in the Chuanjie Formation, Yunnan Province, China. The morphology of the borings closely matches the ichnogenus Cubiculum. Based on the lack of bioglyphs observed in Cubiculum ornatus, a new ichnospecies is proposed here. The new trace fossil, Cubiculum inornatus isp. nov., is interpreted to have been constructed for pupation by an unknown taxon of insect. Additionally, we report even older borings from Early Jurassic dinosaur bones of the Elliott Formation in the Karoo Basin, which represent the second oldest occurrence of insect traces in bone from continental settings. Both trace fossils sites have palaeogeographic implications for the origins and dispersal of osteophagia amongst terrestrial invertebrates during the Mesozoic. These discoveries push back the antiquity of pupation in animal bones by more than 100 million years to the Middle Jurassic, indicating that this behaviour, and osteophagy more generally, originated early in the Mesozoic, roughly comparable with the origination of insect pupation in woody substrates (Late Triassic).     

Insect ovipositions on aquatic plant leaves <Emphasis Type="Italic">Quereuxia</Emphasis> from the Upper Cretaceous of the Amur region

New form taxa of insect ovipositions on aquatic plant leaves Quereuxia from the Campanian locality of Udurchukan (Amur Region) are described. Endophytic ovipositions Paleoovoidus flabellatus sp. nov. and P. arcuatus sp. nov. do not differ in shape from ovipositions of recent damselflies. Exophytic ovipositions Palaexovoidus ovoideus gen. et sp. nov., P. catenulatus sp. nov., P. multus sp. nov., and P. amplus sp. nov. belong to insects that develop in the water, probably dragonflies of the suborder Anisoptera. A new family Palaexovoididae fam. nov. is erected.     

中国中生代昆虫化石研究新进展

回顾了1990年以来中国中生代昆虫化石研究的新进展,简述了我国学者在古昆虫分类学、昆虫内部形态学、昆虫系统学、古生态学、昆虫生物地层学、昆虫区系的历史演变、生物古地理学、昆虫与植物的关系等8个方面取得的突出贡献,并给出了我国发现的世界最低层位昆虫科级类群表。     

中国中生代哺乳类化石

本文对迄今所知的中国中生代哺乳类的化石地点、层位、时代、化石材料和系统分类位置作了总结和讨论。     

The earliest evidence of holometabolan insect pupation in conifer wood

Background

The pre-Jurassic record of terrestrial wood borings is poorly resolved, despite body fossil evidence of insect diversification among xylophilic clades starting in the late Paleozoic. Detailed analysis of borings in petrified wood provides direct evidence of wood utilization by invertebrate animals, which typically comprises feeding behaviors.

Methodology/Principal Findings

We describe a U-shaped boring in petrified wood from the Late Triassic Chinle Formation of southern Utah that demonstrates a strong linkage between insect ontogeny and conifer wood resources. Xylokrypta durossi new ichnogenus and ichnospecies is a large excavation in wood that is backfilled with partially digested xylem, creating a secluded chamber. The tracemaker exited the chamber by way of a small vertical shaft. This sequence of behaviors is most consistent with the entrance of a larva followed by pupal quiescence and adult emergence — hallmarks of holometabolous insect ontogeny. Among the known body fossil record of Triassic insects, cupedid beetles (Coleoptera: Archostemata) are deemed the most plausible tracemakers of Xylokrypta, based on their body size and modern xylobiotic lifestyle.

Conclusions/Significance

This oldest record of pupation in fossil wood provides an alternative interpretation to borings once regarded as evidence for Triassic bees. Instead Xylokrypta suggests that early archostematan beetles were leaders in exploiting wood substrates well before modern clades of xylophages arose in the late Mesozoic.     

Early evolution and historical biogeography of fishflies (Megaloptera: Chauliodinae): implications from a phylogeny combining fossil and extant taxa

Fishflies (Corydalidae: Chauliodinae) are one of the main groups of the basal holometabolous insect order Megaloptera, with ca. 130 species distributed worldwide. A number of genera from the Southern Hemisphere show remarkably disjunctive distributions and are considered to be the austral remnants or "living fossils" of Gondwana. Hitherto, the evolutionary history of fishflies remains largely unexplored due to limited fossil record and incomplete knowledge of phylogenetic relationships. Here we describe two significant fossil species of fishflies, namely Eochauliodes striolatus gen. et sp. nov. and Jurochauliodes ponomarenkoi Wang & Zhang, 2010 (original designation for fossil larvae only), from the Middle Jurassic of Inner Mongolia, China. These fossils represent the earliest fishfly adults. Furthermore, we reconstruct the first phylogenetic hypothesis including all fossil and extant genera worldwide. Three main clades within Chauliodinae are recognized, i.e. the Dysmicohermes clade, the Protochauliodes clade, and the Archichauliodes clade. The phylogenetic and dispersal-vicariance (DIVA) analyses suggest Pangaean origin and global distribution of fishflies before the Middle Jurassic. The generic diversification of fishflies might have happened before the initial split of Pangaea, while some Gondwanan-originated clades were likely to be affected by the sequential breakup of Pangaea. The modern fauna of Asian fishflies were probably derived from their Gondwanan ancestor but not the direct descendents of the Mesozoic genera in Asia.     

The quality of the fossil record of Mesozoic birds

The Mesozoic fossil record has proved critical for understanding the early evolution and subsequent radiation of birds. Little is known, however, about its relative completeness: just how 'good' is the fossil record of birds from the Mesozoic? This question has come to prominence recently in the debate over differences in estimated dates of origin of major clades of birds from molecular and palaeontological data. Using a dataset comprising all known fossil taxa, we present analyses that go some way towards answering this question. Whereas avian diversity remains poorly represented in the Mesozoic, many relatively complete bird specimens have been discovered. New taxa have been added to the phylogenetic tree of basal birds, but its overall shape remains constant, suggesting that the broad outlines of early avian evolution are consistently represented: no stage in the Mesozoic is characterized by an overabundance of scrappy fossils compared with more complete specimens. Examples of Neornithes (modern orders) are known from later stages in the Cretaceous, but their fossils are rarer and scrappier than those of basal bird groups, which we suggest is a biological, rather than a geological, signal.     

Time flies, a new molecular time-scale for brachyceran fly evolution without a clock

The insect order Diptera, the true flies, contains one of the four largest Mesozoic insect radiations within its suborder Brachycera. Estimates of phylogenetic relationships and divergence dates among the major brachyceran lineages have been problematic or vague because of a lack of consistent evidence and the rarity of well-preserved fossils. Here, we combine new evidence from nucleotide sequence data, morphological reinterpretations, and fossils to improve estimates of brachyceran evolutionary relationships and ages. The 28S ribosomal DNA (rDNA) gene was sequenced for a broad diversity of taxa, and the data were combined with recently published morphological scorings for a parsimony-based phylogenetic analysis. The phylogenetic topology inferred from the combined 28S rDNA and morphology data set supports brachyceran monophyly and the monophyly of the four major brachyceran infraorders and suggests relationships largely consistent with previous classifications. Weak support was found for a basal brachyceran clade comprising the infraorders Stratiomyomorpha (soldier flies and relatives), Xylophagomorpha (xylophagid flies), and Tabanomorpha (horse flies, snipe flies, and relatives). This topology and similar alternative arrangements were used to obtain Bayesian estimates of divergence times, both with and without the assumption of a constant evolutionary rate. The estimated times were relatively robust to the choice of prior distributions. Divergence times based on the 28S rDNA and several fossil constraints indicate that the Brachycera originated in the late Triassic or earliest Mesozoic and that all major lower brachyceran fly lineages had near contemporaneous origins in the mid-Jurassic prior to the origin of flowering plants (angiosperms). This study provides increased resolution of brachyceran phylogeny, and our revised estimates of fly ages should improve the temporal context of evolutionary inferences and genomic comparisons between fly model organisms.     

A temperate palaeodiversity peak in Mesozoic dinosaurs and evidence for Late Cretaceous geographical partitioning

Aim Modern biodiversity peaks in the tropics and declines poleward, a pattern that is potentially driven by climate. Although this latitudinal biodiversity gradient (LBG) also characterizes the marine invertebrate fossil record, distributions of ancient terrestrial faunas are poorly understood. This study utilizes data on the dinosaur fossil record to examine spatial patterns in terrestrial biodiversity throughout the Mesozoic. Location We compiled data on fossil occurrences across the globe. Methods We compiled a comprehensive dataset of Mesozoic dinosaur genera (738), including birds. Following the utilization of sampling standardization techniques to mediate for the uneven sampling of the fossil record, we constructed latitudinal patterns of biodiversity from this dataset. Results The dominant group of Mesozoic terrestrial vertebrates did not conform to the modern LBG. Instead, dinosaur diversity was highest at temperate palaeolatitudes throughout the 160 million year span of dinosaurian evolutionary history. Latitudinal diversity correlates strongly with the distribution of land area. Late Cretaceous sauropods and ornithischians exhibit disparate LBGs. Main conclusions The continuity of the palaeotemperate peak in dinosaur diversity indicates a diminished role for climate on the Mesozoic LBG; instead, dinosaur diversity may have been driven by the amount of land area among latitudinal belts. There is no evidence that the tropics acted as a cradle for dinosaur diversity. Geographical partitioning among major clades of herbivorous dinosaurs in the Late Cretaceous may result from the advanced stages of continental fragmentation and/or differing responses to increasing latitudinal climatic zonation. Our results suggest that the modern‐day LBG on land was only established 30 million years ago, following a significant post‐Eocene recalibration, potentially related to increased seasonality.     

The fossil record and taphonomy of butterflies and moths (Insecta,Lepidoptera): implications for evolutionary diversity and divergence-time estimates

Background

It is conventionally accepted that the lepidopteran fossil record is significantly incomplete when compared to the fossil records of other, very diverse, extant insect orders. Such an assumption, however, has been based on cumulative diversity data rather than using alternative statistical approaches from actual specimen counts.

Results

We reviewed documented specimens of the lepidopteran fossil record, currently consisting of 4,593 known specimens that are comprised of 4,262 body fossils and 331 trace fossils. The temporal distribution of the lepidopteran fossil record shows significant bias towards the late Paleocene to middle Eocene time interval. Lepidopteran fossils also record major shifts in preservational style and number of represented localities at the Mesozoic stage and Cenozoic epoch level of temporal resolution. Only 985 of the total known fossil specimens (21.4%) were assigned to 23 of the 40 extant lepidopteran superfamilies. Absolute numbers and proportions of preservation types for identified fossils varied significantly across superfamilies. The secular increase of lepidopteran family-level diversity through geologic time significantly deviates from the general pattern of other hyperdiverse, ordinal-level lineages.

Conclusion

Our statistical analyses of the lepidopteran fossil record show extreme biases in preservation type, age, and taxonomic composition. We highlight the scarcity of identified lepidopteran fossils and provide a correspondence between the latest lepidopteran divergence-time estimates and relevant fossil occurrences at the superfamily level. These findings provide caution in interpreting the lepidopteran fossil record through the modeling of evolutionary diversification and in determination of divergence time estimates.

Electronic supplementary material

The online version of this article (doi:10.1186/s12862-015-0290-8) contains supplementary material, which is available to authorized users.     

Phylogenetic Distribution of Extant Richness Suggests Metamorphosis Is a Key Innovation Driving Diversification in Insects

Insects and their six-legged relatives (Hexapoda) comprise more than half of all described species and dominate terrestrial and freshwater ecosystems. Understanding the macroevolutionary processes generating this richness requires a historical perspective, but the fossil record of hexapods is patchy and incomplete. Dated molecular phylogenies provide an alternative perspective on divergence times and have been combined with birth-death models to infer patterns of diversification across a range of taxonomic groups. Here we generate a dated phylogeny of hexapod families, based on previously published sequence data and literature derived constraints, in order to identify the broad pattern of macroevolutionary changes responsible for the composition of the extant hexapod fauna. The most prominent increase in diversification identified is associated with the origin of complete metamorphosis, confirming this as a key innovation in promoting insect diversity. Subsequent reductions are recovered for several groups previously identified as having a higher fossil diversity during the Mesozoic. In addition, a number of recently derived taxa are found to have radiated following the development of flowering plant (angiosperm) floras during the mid-Cretaceous. These results reveal that the composition of the modern hexapod fauna is a product of a key developmental innovation, combined with multiple and varied evolutionary responses to environmental changes from the mid Cretaceous floral transition onward.     

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.     

New bethylid and chrysidid wasps (Hymenoptera: Chrysidoidea) from Canadian Late Cretaceous amber

A new bethylid species, Celonophamia granama, and two new chrysidid species, Procleptes eoliami, and P. hopejohnsonae, are described from Late Cretaceous (Campanian) amber collected at the Grassy Lake locality in Alberta, Canada. Within the deposit these taxa constitute the first bethylid, and the second and third chrysidid species to be described, respectively. The new taxa expand the sparse fossil record of Chrysidoidea, particularly that of Chrysididae—a group that was previously represented by only three described species in the Mesozoic. The presence of Celonophamia species in both Canadian amber and Siberian (Taimyr) amber further emphasizes faunal similarities between these two northern Late Cretaceous amber deposits. Given the prevalence of metallic coloration in Chrysididae, the specimens described here also provide evidence for the taphonomic alteration of perceived insect colors in Cretaceous amber inclusions.     

中国中生代鞘翅目化石研究现状

1923年A.W.Grabau通过对山东省莱阳盆地莱阳组早白垩世昆虫化石的研究,命名了我国第1种鞘翅目化石昆虫(Proteroscarabaeus yeni).此后,中国昆虫化石分类学者先后发表了40余篇关于我国中生代鞘翅目化石昆虫的分类论著,共描述鉴定了182种,涉及35科,137属.这些化石分布于12个省,其中三叠纪11个种,侏罗纪68个种,白垩纪103个种.本文通过图表列举出我国已发现的中生代鞘翅目化石昆虫名录以及它们的分布和年代,总结了国内中生代甲虫化石当前的研究状况,并初步分析了研究中存在的一些问题.另外本文还介绍了国外学者对于中国古昆虫学者所建立的4个新科提出的不同意见.     

Dinosaurs and fluctuating sea levels during the mesozoic

The very different frequency of dinosaurs during the Mesozoic can be allied to the correlation between global sea level cyclicity and fossilization. This is based upon the sedimentary situation in the inner shelf, the area of predominant fossil record of dinosaurs, and sea level fluctuations. A rich fossil record is found in times of high sea level, and vice versa. Due to natural laws acting on sea level stands, the fossil record of dinosaurs and other terrestrial tetrapods is incomplete. This is causally explainable in the sequence stratigraphy. Among causes of global sea level fluctuations, the change from warm to cold times has been accorded greatest probability even in the Mesozoic. Consequently, the problem of dinosaur evolution and distribution should not be confused with the pattern of their fossil record. The latter, however, is so far nearly always used for all interpretations. The context presented here results in basic modifications.

During the phases of reduced to missing fossil record (low sea level, cold times), dinosaurs existed at least in circumequatorial regions in high diversity. Highly diverse faunas recorded exceptionally in the Upper Jurassic, Middle and Late Cretaceous, were each time the result of a long previous evolution and not the result of short term radiations at these times. Phases of sea level highstand and warm times caused an increased fossil record and poleward distribution. Cretaceous dinosaurs in paleolatitudes of 70° to 80° N and S are no proof for endothermy, but are only the effect of favorable climatic conditions at limited times. Any endothermy of the dinosaurs is not coincident with the supposedly uniformly warm equable climate of the Mesozoic, but with the opposite. Cold times did not hamper the existence of dinosaurs, but led in extreme cases (Aalenian and Valanginian) to the global lack of their fossil record. The situation at the Cretaceous‐Tertiary boundary is also explainable in this context. According to the sea level cyclicity, no extreme sea level fall and no globablly cold time were present in the critical time segment. The regression in the late Maastrichtian is found to belong to a sequence of third‐order cycles beginning in the Campanian. Every one of the cycle boundaries with regression and transgression produced apparent extinction effects which in reality are only gaps in the fossil record. After the late Maastrichtian regression the dinosaurs persisted with six lineages. The so far youngest dinosaur fauna in the Puercan (basal Paleocene) lies in a phase of sea level highstand of minor amplitude and duration with comparatively minor chances for a fossil record. The occurrences in the Puercan are governed by natural law, and, thus, dinosaurs are untied from the short term problems of the Cretaceous‐Tertiary boundary. Why dinosaurs are then missing at the next highstand, remains an open question. Anyhow, mechanisms which control fossil record, diversification and distribution, including global cold periods, do not belong to the direct causes of extinction, because identical occurrences happened many times during the Mesozoic without inducing extinction.     

A new,well-preserved genus and species of fossil Glaphyridae (Coleoptera,Scarabaeoidea) from the Mesozoic Yixian Formation of Inner Mongolia,China

A new genus and species of fossil Glaphyridae, Cretohypna cristata gen. et sp. n., is described and illustrated from the Mesozoic Yixian Formation. This new genus is characterized by the large body; large and strong mandibles; short labrum; elytra without longitudinal carina; and male meso- and possible metatibia apically modified. A list of described fossil glaphyrids of the world is provided. This significant finding broadens the known diversity of Glaphyridae in the Mesozoic China.