The oldest known clerid fossils from the Middle Jurassic of China,with a review of Cleridae systematics (Coleoptera)

Three fossil beetles recorded from Inner Mongolia, China (Callovian, Middle Jurassic, approximately 165 Ma) are assigned to Cleridae and constitute the earliest known representatives of this family. Two of the fossils are described as Protoclerus korynetoides gen. et sp.n. and the third as Wangweiella calloviana gen. et sp.n. Ninety‐six morphological characters of the larvae and adults of 17 extant clerid representatives and seven cleroid families as outgroups were analysed under maximum parsimony with, and without, fossil species included in the matrix. The results indicate that Protoclerus gen.n. is a separate lineage within the family, whereas Wangweiella gen.n. is sister to the rest of Epiclininae. The resulting trees are compared with the most recent morphological and molecular phylogenies of Cleridae. The trees presented appear congruent with the latest molecular phylogenies. The evolution of Cleridae, as well as their biogeography and systematics, are briefly reviewed. A complete list of clerid fossil species described to date is provided. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:32240D3D-2A7A-41F6-B485-A66F2265CB8A .     

中国长翅目昆虫化石研究现状

长翅目昆虫在地史纪录上可以追溯到早二叠纪,是全变态昆虫中最原始的种类之一.截至目前,29篇关于中国长翅目昆虫化石分类的论著发表,共描述我国长翅目昆虫化石11科28属51种,这些化石分布于从三叠纪到白垩纪的不同地层中.本文通过图表提供了我国已发现的长翅目化石名录并介绍了其分布和年代,回顾了我国长翅目昆虫化石的研究进展,指出了一些分类存在的问题,简要慨述了长翅目中一些科的起源与演化以及长翅目与其它全变态昆虫的关系.我国长翅目化石种类丰富,但在基础分类、系统演化方面还缺乏系统的研究,因而许多工作甚至最基础的分类工作亟待开展.     

Discovery of Aspidytidae,a new family of aquatic Coleoptera

The six extant aquatic families of Hydradephaga (Coleoptera) known so far represent a diverse group of beetles morphologically highly modified for life in the water. We report the discovery of a new genus with two species from South Africa and China, which differ greatly from all extant families, but resemble the Jurassic-Cretaceous dagger Liadytidae (the dagger symbol indicates that the taxa are known only as fossils). Based on a combined phylogenetic analysis of molecular and morphological data we erect a new family, Aspidytidae, which is the sister group of Dytiscidae plus Hygrobiidae. We propose a new scenario for the evolution of swimming behaviour in adephagan beetles, in which the transition into the aquatic environment is followed by complex and repeated changes in lifestyles, including the secondary complete loss of swimming ability in Aspidytidae.     

松科早期球果研究的历史和现状

本文主要对早白垩世到老第三纪渐新世这段时间的松科化石球果的研究作了系统回顾和总结。松科化石球果的研究经历了一个由分散、幼稚到系统、成熟的发展过程,现在将松科化石球果分为三个属：Pityostrobus、Pseudoaraucaria 及Pinus。其中Pseudoaraucaria及Pinus为两个自然属,Pseudoaraucaria已经绝灭。Pityostrobus作为一个器官属,包括那些在松科中分类位置不明的球果。对松科化石球果进行细致的外部形态及内部结构的研究主要目的在于演化关系的探讨上：对于Pinus,主要探讨这个属内各个亚属、组、亚组之间的演化关系;对于Pityostrobus和Pseudoaraucaria,通过绝灭类型的分析,探索松科内各个类群(包括绝灭的)的演化关系进而完成进化树是研究的主要方向,现在这方面已有了Pinus中心说及Pseudoaraucaria分支说两个假说。     

A review of Quaternary range shifts in European aquatic Coleoptera

Aim To undertake a quantitative review of the Quaternary fossil record of European water beetles to evaluate their geographical and temporal coverage, and to characterize the extent and typology of the shifts in their geographical ranges. Location Europe. Methods We compiled Quaternary water beetle records from public databases and published references. We included in the analyses species of 10 families of aquatic Coleoptera, and recorded range shifts through the comparison of the location of fossil remains with the current distribution of the species. We explored the ecological representativeness of the fossil record, as well as the relationship between range shifts and the habitat type of the species. Results Our final data set included over 9000 records for 259 water beetle species. Fossil remains of aquatic beetles have been documented exclusively north of 42° N, with most of the records from the British Isles and virtually none from southern Europe or the Mediterranean Basin. Over 80% of the records were from the Late Glacial and the Holocene periods (the last 15 kyr), and overall approximately 20% of the species have been recorded outside their present range (23% excluding Holocene records). Most range shifts were southern or western extensions of currently widespread, northern species, with 10 species displaying major range shifts through the Palaearctic. Lentic species were significantly more likely to have experienced major range shifts, even accounting for the general ecological bias of the fossil record towards lentic habitats. Main conclusions Our results show that the Quaternary record of aquatic Coleoptera is geographically, temporally and ecologically skewed, necessitating caution when extrapolating general conclusions about range changes and ecological stability to other areas or periods on the basis of such scattered evidence. Most central and northern European species for which there are fossil records seem to have conserved their ranges through the Late Pleistocene, with geographical shifts mostly restricted to species with current widespread north Palaearctic or Holarctic distributions. Major range shifts through the Palaearctic are taxonomically uneven, suggesting either an idiosyncratic behaviour of taxa depending on ecological or phylogenetic factors, or a sampling artefact produced by the limited availability of taxonomic expertise.     

INTEGRATING FOSSILS WITH MOLECULAR PHYLOGENIES IMPROVES INFERENCE OF TRAIT EVOLUTION

Comparative biologists often attempt to draw inferences about tempo and mode in evolution by comparing the fit of evolutionary models to phylogenetic comparative data consisting of a molecular phylogeny with branch lengths and trait measurements from extant taxa. These kinds of approaches ignore historical evidence for evolutionary pattern and process contained in the fossil record. In this article, we show through simulation that incorporation of fossil information dramatically improves our ability to distinguish among models of quantitative trait evolution using comparative data. We further suggest a novel Bayesian approach that allows fossil information to be integrated even when explicit phylogenetic hypotheses are lacking for extinct representatives of extant clades. By applying this approach to a comparative dataset comprising body sizes for caniform carnivorans, we show that incorporation of fossil information not only improves ancestral state estimates relative to those derived from extant taxa alone, but also results in preference of a model of evolution with trend toward large body size over alternative models such as Brownian motion or Ornstein–Uhlenbeck processes. Our approach highlights the importance of considering fossil information when making macroevolutionary inference, and provides a way to integrate the kind of sparse fossil information that is available to most evolutionary biologists.     

On the head morphology of Tetraphalerus, the phylogeny of Archostemata and the basal branching events in Coleoptera

Internal and external features of Tetraphalerus bruchi were studied using X‐ray microtomography (µ‐CT) and other techniques, and head structures were described in detail. µ‐Ct is highly efficient for the assessment of anatomical data. A data matrix with 90 morphological characters of recent and fossil beetles was analyzed with different approaches (parsimony, Bayesian analysis). The results of the parsimony analysis resulted in the following branching pattern: (?Tshekardocoleidae + (?Permocupedidae, ?Rhombocoleidae + (?Triadocupedidae + ((Adephaga + (Myxophaga + Polyphaga))) + Archostemata s.str. [including Jurodidae]))). Sikhotealinia is placed as sister group of ?Jurodes (Jurodidae), and Jurodidae as sister group of the remaining Archostemata (Bayesian analysis) or of a clade comprising Micromalthidae, Crowsoniellidae, ?Ademosynidae, ?Schizophoridae and ?Catiniidae. The monophyly of Ommatidae and Cupedidae is well supported and Priacma is placed as the sister group of all other Cupedidae. Important events in the early evolution of Coleoptera are the shortening of the elytra and the transformation of the elytral venation (Coleoptera excluding ?Tshekardocoleidae), the formation of a closed subelytral space (Coleoptera excluding ?Tshekardocoleidae and ?Permocupedidae), the reduction of two apical antennomeres, and the loss of the broad prothoracic postcoxal bridge (Coleoptera excluding ?Tshekardocoleidae, ?Permocupedidae and ?Rhombocoleidae). Plesiomorphic features preserved in extant Archostemata are the tuberculate cuticle, the elytral pattern with parallel longitudinal ribs and window punctures, a mesoventrite with a transverse ridge, triangular mesocoxae with a distinct meron, and the exposed metatrochantin. The fossils included in the analyses do not only contribute to the reconstruction of character evolution but also influence the branching pattern. An understanding of the major evolutionary events in Coleoptera would not be possible without considering the rich fossil record of Permian and Mesozoic beetles. © The Willi Hennig Society 2007.     

Microscopic,chemical and molecular methods for examining fossil preservation

Advances in technology over the past two decades have resulted in unprecedented access to data from biological specimens. These data have expanded our understanding of physical characteristics, physiological, cellular and subcellular processes, and evolutionary relationships at the molecular level and beyond. Paleontological and archaeological sciences have recently begun to apply these technologies to fossil and subfossil representatives of extinct organisms. Data derived from multidisciplinary, non-traditional techniques can be difficult to decipher, and without a basic understanding of the type of information provided by these methods, their usefulness for fossil studies may be overlooked. This review describes some of these powerful new analytical tools, the data that may be accessible through their use, advantages and limitations, and how they can be applied to fossil material to elucidate characteristics of extinct organisms and their paleoecological environments.     

Ancestral state reconstruction of body size in the Caniformia (Carnivora, Mammalia): the effects of incorporating data from the fossil record

A recent molecular phylogeny of the mammalian order Carnivora implied large body size as the ancestral condition for the caniform subclade Arctoidea using the distribution of species mean body sizes among living taxa. "Extant taxa-only" approaches such as these discount character state observations for fossil members of living clades and completely ignore data from extinct lineages. To more rigorously reconstruct body sizes of ancestral forms within the Caniformia, body size and first appearance data were collected for 149 extant and 367 extinct taxa. Body sizes were reconstructed for four ancestral nodes using weighted squared-change parsimony on log-transformed body mass data. Reconstructions based on extant taxa alone favored large body sizes (on the order of 10 to 50 kg) for the last common ancestors of both the Caniformia and Arctoidea. In contrast, reconstructions incorporating fossil data support small body sizes (< 5 kg) for the ancestors of those clades. When the temporal information associated with fossil data was discarded, body size reconstructions became ambiguous, demonstrating that incorporating both character state and temporal information from fossil taxa unambiguously supports a small ancestral body size, thereby falsifying hypotheses derived from extant taxa alone. Body size reconstructions for Caniformia, Arctoidea, and Musteloidea were not sensitive to potential errors introduced by uncertainty in the position of extinct lineages relative to the molecular topology, or to missing body size data for extinct members of an entire major clade (the aquatic Pinnipedia). Incorporating character state observations and temporal information from the fossil record into hypothesis testing has a significant impact on the ability to reconstruct ancestral characters and constrains the range of potential hypotheses of character evolution. Fossil data here provide the evidence to reliably document trends of both increasing and decreasing body size in several caniform clades. More generally, including fossils in such analyses incorporates evidence of directional trends, thereby yielding more reliable ancestral character state reconstructions.     

松科早期球果研究的历史与现状

本文主要对早白垩世到老第三纪渐新世这段时间的松科化石球果的研究作了系统回顾和总结。松科化石球果的研究经历了一个由分散、幼稚到系统、成熟的发展过程,现在将松科化石球果分为三个属：Ｐｉｔｙｏｓｔｒｏｂｕｓ、Ｐｓｅｕｄｏａｒａｕｃａｒｉａ及Ｐｉｎｕｓ。其中Ｐｓｅｕｄｏａｒａｕｃａｒｉａ及Ｐｉｎｕｓ为两个自然属,Ｐｓｅｕｄｏａｒａｕｃａｒｉａ已经绝灭。Ｐｉｔｙｏｓｔｒｏｂｕｓ作为一个器官属,包括那些在松科中分类位置不明的球果。对松科化石球果进行细致的外部形态及内部结构的研究主要目的在于演化关系的探讨上：对于Ｐｉｎｕｓ,主要探讨这个属内各个亚属、组、亚组之间的演化关系;对于Ｐｉｔｙｏｓｔｒｏｂｕｓ和Ｐｓｅｕｄｏａｒａｕｃａｒｉａ,通过绝灭类型的分析,探索松科内各个类群（包括绝灭的）的演化关系进而完成进化树是研究的主要方向,现在这方面已有了Ｐｉｎｕｓ中心说及Ｐｓｅｕｄｏａｒａｕｃａｒｉａ分支说两个假说。     

水生甲虫系统发育研究进展

简要回顾了水生甲虫近十几年来各个分类阶元水平系统关系的研究历史,对代表性的水生甲虫类高级阶元系统发育关系进行了比较分析,总结了普遍存在的共识和争议.在研究进展中还介绍了分子生物学方法在水生甲虫系统发育研究中的尝试,虽然其研究结果还未能完全解决上述问题,但必将是对现有研究方法的补充.     

Molecular evidence of cycad seed predation by immature Aulacoscelidinae (Coleoptera: Orsodacnidae)

Adult beetles in the small subfamily Aulacoscelidinae (superfamily Chrysomeloidea) are known to feed on the foliage and juices of New World cycads (Order Cycadales; family Zamiaceae), but the habits of larvae have long remained a mystery. We provide the first direct evidence that Aulacoscelidinae larvae feed on and develop within the megagametophyte of the Mesoamerican cycad, Dioon merolae (Zamiaceae). Phylogenetic analyses based on partial DNA sequences from 3 genes recover a cycad seed‐feeding larva proposed to belong to Aulacoscelidinae. These observations reveal a more intimate feeding relationship between Aulacoscelidinae and their New World cycad host plants than was previously recognized. Further, adult Aulacoscelidinae have long been noted to resemble Jurassic fossil chrysomeloids in the extinct subfamily Protoscelidinae. The molecular, morphological, ecological and fossil data reported herein are broadly compatible with an early association between Aulacoscelidinae and their gymnosperm hosts.     

Relationship between osteology and aquatic locomotion in birds: determining modes of locomotion in extinct Ornithurae

The evolutionary history of aquatic invasion in birds would be incomplete without incorporation of extinct species. We show that aquatic affinities in fossil birds can be inferred by multivariate analysis of skeletal features and locomotion of 245 species of extant birds. Regularized discriminant analyses revealed that measurements of appendicular skeletons successfully separated diving birds from surface swimmers and flyers, while also discriminating among different underwater modes of swimming. The high accuracy of this method allows detection of skeletal characteristics that are indicative of aquatic locomotion and inference of such locomotion in bird species with insufficient behavioural information. Statistical predictions based on the analyses confirm qualitative assessments for both foot‐propelled (Hesperornithiformes) and wing‐propelled (Copepteryx) underwater locomotion in fossil birds. This is the first quantitative inference of underwater modes of swimming in fossil birds, enabling future studies of locomotion in extinct birds and evolutionary transitions among locomotor modes in avian lineage.     

Evolution of fossil and living spider flies based on morphological and molecular data (Diptera,Acroceridae)

The phylogeny of spider flies is presented based on an analysis of DNA sequence data combined with morphological characters for both living and fossil species. We sampled 40 extant and extinct genera across all major lineages of Acroceridae, which were compared with outgroup taxa from various lower brachyceran families. In all, 81 morphological characters of 60 extant and 10 extinct ingroup species were combined with 7.1 kb of DNA sequences of two nuclear (CAD and 28S rDNA) and two mitochondrial genes (COI and 16S rDNA). Results strongly support the monophyly of Acroceridae, with major clades contained within classified here in five extant subfamilies (Acrocerinae, Cyrtinae stat. rev. , Ogcodinae stat. rev. , Panopinae and Philopotinae) and one extinct subfamily, Archocyrtinae. The evolution of important spider fly traits is discussed, including genitalia and wing venation. The status of the enigmatic Psilodera Gray and Pterodontia Gray as members of the Panopinae is confirmed based on both molecular and morphological data.     

中生代鞘翅目扁甲总科化石的研究进展

扁甲总科隶属于鞘翅目多食亚目扁甲系,目前包含25个现生科和3个化石科,其中10个科仅分布于南半球,特别是澳大利亚、新西兰和南美。当前的分子系统发育学研究显示扁甲总科的起源时间为晚三叠世至早侏罗世,而冈瓦纳古陆与劳亚古陆的分离也发生在早侏罗世约1.8亿年前。因此对中生代扁甲总科化石的研究是解释扁甲总科如今分布格局的重要基础,也有助于研究扁甲总科的演化历史。本文简要回顾了现生扁甲总科的研究历史,并对其中生代的化石研究进行了概述,迄今在中生代地层发现有12科54种扁甲总科化石。     

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.     

Diversity of aquatic and riparian Coleoptera of the Philippines: checklist,state of knowledge,priorities for future research and conservation

The diversity of the aquatic and strictly riparian beetles of the Philippines is reviewed. A checklist (with 317 species and subspecies) of the aquatic and riparian Coleoptera of the Philippines is provided for the first time herein. This checklist covers the following families (number of species and subspecies in parentheses): Hydroscaphidae (1), Gyrinidae (15), Dytiscidae (65), Noteridae (4), Haliplidae (2), Hydraenidae (15), Hydrochidae (2), Spercheidae (1), Hydrophilidae (72), Scirtidae (49), Elmidae (23), Dryopidae (2), Psephenidae (16), Eulichadidae (1), Limnichidae (20), Heteroceridae (5), Malachiidae (5), Lampyridae (15), Nitidulidae (4). A total of 201 species and subspecies (63%) are considered to be endemic to the Philippines. Aquatic/riparian beetle genera and families that are still unrecorded but likely to occur in this country are briefly discussed. Based on estimates at the genus level, it is concluded that about 850 aquatic and riparian species can be expected to occur in the Philippines. The implication of the Philippine biogeographic history for this diversity is discussed. The potential of aquatic/riparian beetles as bioindicators is briefly examined. The dilemma of aquatic invertebrate conservation is analysed and priorities for future research are outlined.