Wing Shape of Four New Bee Fossils (Hymenoptera: Anthophila) Provides Insights to Bee Evolution

Bees (Anthophila) are one of the major groups of angiosperm-pollinating insects and accordingly are widely studied in both basic and applied research, for which it is essential to have a clear understanding of their phylogeny, and evolutionary history. Direct evidence of bee evolutionary history has been hindered by a dearth of available fossils needed to determine the timing and tempo of their diversification, as well as episodes of extinction. Here we describe four new compression fossils of bees from three different deposits (Miocene of la Cerdanya, Spain; Oligocene of Céreste, France; and Eocene of the Green River Formation, U.S.A.). We assess the similarity of the forewing shape of the new fossils with extant and fossil taxa using geometric morphometrics analyses. Predictive discriminant analyses show that three fossils share similar forewing shapes with the Apidae [one of uncertain tribal placement and perhaps near Euglossini, one definitive bumble bee (Bombini), and one digger bee (Anthophorini)], while one fossil is more similar to the Andrenidae. The corbiculate fossils are described as Euglossopteryx biesmeijeri De Meulemeester, Michez, & Engel, gen. nov. sp. nov. (type species of Euglossopteryx Dehon & Engel, n. gen.) and Bombus cerdanyensis Dehon, De Meulemeester, & Engel, sp. nov. They provide new information on the distribution and timing of particular corbiculate groups, most notably the extension into North America of possible Eocene-Oligocene cooling-induced extinctions. Protohabropoda pauli De Meulemeester & Michez, gen. nov. sp. nov. (type species of Protohabropoda Dehon & Engel, n. gen.) reinforces previous hypotheses of anthophorine evolution in terms of ecological shifts by the Oligocene from tropical to mesic or xeric habitats. Lastly, a new fossil of the Andreninae, Andrena antoinei Michez & De Meulemeester, sp. nov., further documents the presence of the today widespread genus Andrena Fabricius in the Late Oligocene of France.     

New fossil Ochteridae (Hemiptera: Heteroptera: Ochteroidea) from the Upper Mesozoic of north‐eastern China,with phylogeny of the family

Specimens of the velvety shore bugs (Hemiptera: Ochtheridae) occur in the Early Cretaceous Yixian Formation in north‐eastern China. We describe two new genera and three new species –Pristinochterus ovatus Yao, Zhang & Ren sp. nov., Floricaudus multilocellus Yao, Ren & Shih gen.n. et sp.n. and Angulochterus quatrimaculatus Yao, Zhang & Ren gen.n. et sp.n. – from this deposit. A cladistic analysis based on a combination of fossil and extant taxa, and their morphological characters, clarifies the phylogenetic status of the new fossils and allows the reconstruction of relationships within the family Ochteridae. Two main clades within Ochteridae are recognized from the cladistic analysis: Pristinochterus Yao Cai & Ren and Floricaudus Yao, Ren & Shih gen.n. form a monophyletic lineage; and Angulochterus Yao, Zhang & Ren gen.n. emerges as a sister group to all extant velvety shore bugs. A key to all fossil and extant genera of Ochteridae is provided.     

Wing morphology of a new Cretaceous praying mantis solves the phylogenetic jigsaw of early‐diverging extant lineages

The extremely derived morphology and behaviour of extant praying mantises combined with a scarce record of fossil relatives introduce significant challenges to tracing their evolution from Palaeozoic stem‐dictyopterans. Extant members of Chaeteessidae, Mantoididae and Metallyticidae could be invaluable to resolving the mantodean tree, yet their inclusion in phylogenetic analyses led to conflicting hypotheses due to their highly disparate respective morphologies. In this contribution, we present Labradormantis guilbaulti gen. et sp.n. , a new fossil species described from both fore‐ and hind‐wing imprints discovered in the Redmond Mine locality (Late Cretaceous, Cenomanian, Redmond Formation; Labrador, Canada). The examination of its hind‐wing AA2* supports the hypothesis that this structure, unique to Chaeteessidae among extant mantises, is a true vein and that its occurrence represents a plesiomorphy for Mantodea. A parsimony analysis including newly coded wing‐related characters further established that L. guilbaulti gen. et sp.n. displays a unique combination of plesiomorphic and apomorphic character states that situates it within the extinct family Baissomantidae. This dataset resolved the phylogenetic relationships of early‐diverging extant lineages as (Chaeteessidae (Mantoididae (Metallyticidae, Artimantodea))), and suggested that the Eocene Lithophotina floccosa Cockerell might be a close relative of extant metallyticids. It also indicated a trend towards increased modularity within mantis fore‐wings, in contrast with a trend towards increased morphological integration in their hind‐wings, both of which are potentially associated with improved flight performance for modern mantises. This study emphasizes the importance of fossils for resolving phylogenetic relationships and for introducing transitional phenotypes to infer ancient evolutionary trends of extant derived clades.     

New spiders in Upper Cretaceous amber from New Jersey in the American Museum of Natural History (Arthropoda: Araneae)

The oldest described fossils of the extant spider family Araneidae (Araneinae; gen. et sp. indet.), the extant genus Orchestina (Oonopidae; O. sp. indet.) and the new fossil genus Palaeosegestria (Segestriidae; P. lutzzii gen. et sp. nov.) are presented from Upper Cretaceous amber of New Jersey. The known fossil range of the extant family Araneidae is extended approximately 50 myr from the previously oldest described araneid from the Middle Eocene oil shales of the Messel pit in Hesse, Germany. The fossil range of the extant genus Orchestina is also extended 50 myr from the previously oldest described specimen in Eocene Baltic amber.     

Morphological conservatism in the foreleg structure of cicada hatchlings,Burmacicada protera n. gen., n. sp. in Burmese amber,Dominicicada youngi n. gen., n. sp. in Dominican amber and the extant Magicicada septendecim (L.) (Hemiptera: Cicadidae)

Two cicada hatchlings (Hemiptera: Cicadidae) in Burmese and Dominican amber are described as Burmacicada protera n. gen., n. sp. and Dominicicada youngi n. gen., n. sp., respectively. Although very similar in appearance, the two species can be separated by body contour, the nature of the process on the terminal antennomere and the shape and size of protrusions, teeth and spines on the forelegs. A comparison of the forelegs of the fossil hatchlings with those of an extant hatchling of the periodical cicada, Magicicada septendecim (L.), reveals a remarkable degree of morphological conservatism over 100 million years. A brief review of fossil cicadas is presented.     

New braconid wasps from French Cretaceous amber (Hymenoptera, Braconidae): synonymization with Eoichneumonidae and implications for the phylogeny of Ichneumonoidea

Two new fossils of Braconidae are described from Albian-Cenomanian amber of south-western France, Protorhyssalodes arnaudi gen. n., sp. n., and Aenigmabracon capdoliensis gen. n., sp. n. The former appears superficially similar to the type genus and species of the extinct sub-family Protorhyssalinae, from Turonian New Jersey amber specimens, and the latter both to Protorhyssalus and to members of the extinct family Eoichneumonidae. However, both new taxa display unique combinations of wing venation characters making confident assignment to sub-family impossible. Indeed, they are the first braconids ever known to possess both vein 2-CU and a distinct trace of vein 2-1A on hindwing. The new fossil taxa are incorporated into a morphological analysis of extinct and extant ichneumonoids. As a result of the analyses we synonymize the Eoichneumonidae with the Braconidae.     

Geometric morphometric analysis of a new Miocene bumble bee from the Randeck Maar of southwestern Germany (Hymenoptera: Apidae)

The first fossil bumble bee (Apinae: Bombini) from the Miocene Randeck Maar of southwestern Germany is described and illustrated. The specimen is subjected to a geometric morphometric analysis along with a diversity of other bumble bee species representing most major extant lineages, and particularly the subgenus Bombus s.s. The morphometric analysis supports the placement of the Randeck Maar species within Bombus s.s., as a species distinct from all others in the subgenus. It shows that extant subgenera of bumblebees were already derived in the early/middle Miocene. The Randeck Maar fossil is formally described as Bombus (Bombus) randeckensis Wappler & Engel sp. n. .     

Eocene tortoise beetles from the Green River Formation in Colorado,U.S.A. (Coleoptera: Chrysomelidae: Cassidinae)

Abstract The fossil history of leaf beetles (Chrysomelidae) is relatively poorly documented despite an abundance of available material. Of particular interest is the origin and radiation of the diverse tortoise beetles, a derived group within Cassidinae s.l. (=Cassidinae + Hispinae) defined by the exophagous life history and specialized morphology of the immature stages. Cassidinae is also a group with relatively few fossil records that can be assigned with any degree of certainty. Here we report two of the oldest definitive tortoise beetle fossils, Eosacantha delocranioides gen.n. et sp.n. and Denaeaspis chelonopsis gen.n. et sp.n. , from the Eocene Green River Formation (ca. 47 million years old) in northwestern Colorado, U.S.A. Owing to the fine level of preservation, many important features can be observed and are coded into the recent cladistic analysis for the subfamily. Phylogenetic analysis highlights that both genera have affinities with modern lineages, one restricted to the Old World and the other restricted to the Neotropics. Although Cassidinae as a whole extend into the Cretaceous, the available information suggests that the tortoise beetles perhaps originated and diversified during the Early Tertiary. As such, the morphological and biological transitions from the leaf‐mining hispiforms to the distinctive tortoise‐like cassidiforms, with their elaborate defensive larval shields and other unique behaviours, probably took place during the latest Paleocene or earliest Eocene. These Green River fossils are the oldest yet to document the specialized morphology associated with the transition in cassidine feeding and immature biology.     

On the edge of parasitoidism: a new Lower Cretaceous woodwasp forming the putative sister group of Xiphydriidae + Euhymenoptera

The fossil woodwasp Cratoenigma articulata gen. et sp.n. (Insecta: Hymenoptera) is described from the Lower Cretaceous Crato formation of Brazil. This fossil cannot be placed in any existing superfamily, but its putative phylogenetic position within Hymenoptera is discussed in detail on the basis of relevant thoracic, abdominal and wing venation characters. These characters are critically evaluated and compared with those of extant and fossil Hymenoptera. The phylogenetic position of C. articulata sp.n. is investigated relative to extant Xyelidae, Tenthredinoidea s.l., Pamphilioidea, Cephidae, Siricoidea, Xiphydriidae, Orussidae and Apocrita, and also to Mesozoic Gigasiricidae, Myrmiciidae, Daohugoidae, Sepulcidae, Anaxyelidae, Paroryssidae and Ephialtitidae. Based on the presence of a synapomorphic transscutal articulation, a plesiomorphic unconcealed mesopostnotum and autapomorphic hindwing venation (cu‐a distinctly basal to fork between M and Cu), C. articulata sp.n. most likely forms the sister group of Xiphydriidae + Euhymenoptera. This would place it well within Unicalcarida, i.e. the clade in which the transition from endophytic to parasitoid lifestyle evolved.     

New fossil mousebird (Aves: Coliiformes) with feather preservation provides insight into the ecological diversity of an Eocene North American avifauna

Coliiformes (mousebirds) are represented by just six extant species. These species, restricted to sub‐Saharan Africa, are all primarily frugivorous and are among the most sedentary of living birds. Previously described fossil Coliiformes preserving feather traces share the short, rounded wing shape of extant mousebirds. Along with osteological evidence, these observations have been proposed to support poor sustained flight capabilities across the stem mousebird lineage. We report a new species of Coliiformes from the early Eocene (51.66 ± 0.09 Ma) Fossil Butte Member of the Green River Formation, represented by one of the comparatively few fossils from these deposits preserving carbonized traces of the wing and tail feathering. Feather traces indicate an elongate, tapering wing shape similar to that of some extant aerial insectivores, and suggestive of a capacity for sustained and agile open‐air flight. Traces of the rectrices reveal the tail accounted for approximately two‐thirds of the total length of the bird, a proportion similar to that in living mousebirds. Phylogenetic analysis places the new species as a stem representative of Coliiformes, demonstrating for the first time that the two major clades of Coliiformes – Sandcoleidae and Colii – co‐occurred at Fossil Lake. Based on the recovered phylogeny, as well as the osteology and feathering of extant and fossil Coliiformes, the wing shape of the new species is interpreted as apomorphic. In addition to documenting unexpected morphological specialization within stem‐lineage Coliiformes, the new species adds yet another taxon to the emerging reconstruction of the diverse Paleogene avifauna from the tightly dated and nearly synchronous fossiliferous deposits of the Fossil Butte Member. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 685–706.     

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 .     

A new interpretation of the bee fossil Melitta willardi Cockerell (Hymenoptera,Melittidae) based on geometric morphometrics of the wing

Although bees are one of the major lineages of pollinators and are today quite diverse, few well-preserved fossils are available from which to establish the tempo of their diversification/extinction since the Early Cretaceous. Here we present a reassessment of the taxonomic affinities of Melitta willardi Cockerell 1909, preserved as a compression fossil from the Florissant shales of Colorado, USA. Based on geometric morphometric wing shape analyses M. willardi cannot be confidently assigned to the genus Melitta Kirby (Anthophila, Melittidae). Instead, the species exhibits phenotypic affinity with the subfamily Andreninae (Anthophila, Andrenidae), but does not appear to belong to any of the known genera therein. Accordingly, we describe a new genus, Andrenopteryx gen. n., based on wing shape as well as additional morphological features and to accommodate M. willardi. The new combination Andrenopteryx willardi (Cockerell) is established.     

A new genus and species of scorpionfly (Mecoptera) from Baltic amber,with an unusually developed postnotal organ

A new genus and species of fossil scorpionflies (Mecoptera) Baltipanorpa damzeni gen. et sp.n. is described from two well‐preserved male specimens in Baltic amber (middle Eocene: Lutetian). The most characteristic feature of the new taxon is an unusually developed postnotal organ on abdominal tergum IV. This is the most extremely developed example of this organ among Mecoptera and the only observation of notal and postnotal organs among fossil scorpionflies. The following combination of characters are provided to distinguish the new genus from other Panorpidae: Sc, short; R₁ and R₂ two‐branched; A₁ joins posterior margin of wing only at same level as fork of vein Rs; unusual shape of abdomen, abdominal segments I–IV strongly reduced, abdominal segment V elongate and widened, segments VII and VIII strongly elongate; notal and postnotal organs present, strongly developed process (postnotal organ) on tergum IV, unknown in all described extant and fossil scorpionflies. Different types of notal organs of Mecoptera are compared and their function and morphology are discussed. Morphological analysis of notal and postnotal organs in extant species permits us to conclude that B. damzeni sp.n. is characterized by the most developed and complex notal organs in all Mecoptera.     

Bees diversified in the age of eudicots

Reliable estimates on the ages of the major bee clades are needed to further understand the evolutionary history of bees and their close association with flowering plants. Divergence times have been estimated for a few groups of bees, but no study has yet provided estimates for all major bee lineages. To date the origin of bees and their major clades, we first perform a phylogenetic analysis of bees including representatives from every extant family, subfamily and almost all tribes, using sequence data from seven genes. We then use this phylogeny to place 14 time calibration points based on information from the fossil record for an uncorrelated relaxed clock divergence time analysis taking into account uncertainties in phylogenetic relationships and the fossil record. We explore the effect of placing a hard upper age bound near the root of the tree and the effect of different topologies on our divergence time estimates. We estimate that crown bees originated approximately 123 Ma (million years ago) (113–132 Ma), concurrently with the origin or diversification of the eudicots, a group comprising 75 per cent of angiosperm species. All of the major bee clades are estimated to have originated during the Middle to Late Cretaceous, which is when angiosperms became the dominant group of land plants.     

The oldest bee fly in the French Paleocene (Diptera: Bombyliidae)

Paleolomatia menatensis gen. and sp. n., oldest Bombyliidae sensu stricto, is described from the Paleocene of Menat (France). The new genus, based on the wing venation, is attributed to the rather ‘derived’ subfamily Lomatiinae, strongly supporting a Late Cretaceous age for the diversification of the pollinator bee flies, in relation with the floristic changes and the angiosperm radiation that occurred at the same time.     

A New Snake Skull from the Paleocene of Bolivia Sheds Light on the Evolution of Macrostomatans

Macrostomatan snakes, one of the most diverse extant clades of squamates, display an impressive arsenal of cranial features to consume a vast array of preys. In the absence of indisputable fossil representatives of this clade with well-preserved skulls, the mode and timing of these extraordinary morphological novelties remain obscure. Here, we report the discovery of Kataria anisodonta n. gen. n. sp., a macrostomatan snake recovered in the Early Palaeocene locality of Tiupampa, Bolivia. The holotype consists of a partial, minute skull that exhibits a combination of booid and caenophidian characters, being the presence of an anisodont dentition and diastema in the maxilla the most distinctive trait. Phylogenetic analysis places Kataria basal to the Caenophidia+Tropidophiidae, and represents along with bolyeriids a distinctive clade of derived macrostomatans. The discovery of Kataria highlights the morphological diversity in the maxilla among derived macrostomatans, demonstrating the relevance of maxillary transformations in the evolution of this clade. Kataria represents the oldest macrostomatan skull recovered, revealing that the diversification of macrostomatans was well under way in early Tertiary times. This record also reinforces the importance of Gondwanan territories in the history of snakes, not only in the origin of the entire group but also in the evolution of ingroup clades.     

缅甸克钦琥珀白垩纪石蝇(襀翅目: 襀科)新属种记述

文中简述了襀翅目缅甸琥珀研究概况,并基于一件保存较好的雌虫标本,报道了襀翅目襀科新属种Burperla decolorata gen. et sp. nov.。该化石具有以下重要特征:体较长,体色浅,触须与触角较长, RP脉末端接近翅端部,下殖板宽大、圆形,后缘中部突出。这些形态特征显著区别于襀科的其它化石及现生类群。     

Early Cretaceous origin of pollen‐feeding beetles (Insecta: Coleoptera: Oedemeridae)

The taxonomic position of a new pollen‐feeding fossil beetle from Spanish amber (late Albian, 105 Ma) is analysed. A phylogenetic analysis allows me to accommodate Darwinylus marcosi gen. et sp. nov. in the Polyphaga: Oedemeridae within current limits for the family, which clearly belongs in the subfamily Oedemerinae. It corresponds to the oldest definitive record for the family. Some autapomorphies, mainly in antennae, are observable in the fossil compared with extant members of the family. A discussion about these problematic characters and the evolution of the family is proposed.