Cretaceous and Eocene fossils of the rare extant genus Synneuron Lundstrom (Diptera: Canthyloscledidae): evidence of a true Pangean clade

The first two fossil species of the canthyloscelid genus Synneuron are described based on compression wings. Synneuron eomontana sp. nov. is described from the Middle Eocene Coal Creek Member of the Kishenehn Formation, in the USA, and Synneuron jelli sp. nov. is described from the Lower Cretaceous Koonwarra Fossil Bed of the Korumburra Group, in Australia. The wings are illustrated and compared to the extant species of the genus, to species of the three other recent genera of Canthyloscelidae and to an anisopodid. A phylogenetic analysis of the relationships between the species of Synneuron was performed. The Eocene fossil S. eomontana appears as sister of the pair of recent Holarctic species of the genus, while the Australian Cretaceous species S. jelli is sister of the clade with the species of Synneuron of the northern hemisphere. The sister group of Synneuron is the canthyloscelid clade (Hyperoscelis + Canthyloscelis), for which a middle Jurassic fossil is known. At the early Cretaceous, Gondwana was already separated from Laurasia and the disjunction between the species of Synneuron in Australia and the northern hemisphere clade of the genus suggest a true pangeic origin for the genus. The biology of the canthyloscelid larvae is shaped by its trophic specialization—xylosaprophagous. This suggests that the transition from the Pangean Jurassic gymnosperm-dominated forests to the late Cretaceous angiosperm-dominated forests may be related to the low recent diversity of Synneuron or of the canthyloscelids in the world—and maybe to the extinction of the genus in the southern hemisphere. This major turnover of the vegetation type along the Cretaceous may be also somehow related to the complete extinction of other groups of flies strictly associated with gymnosperms, as may be the case of the lower brachyceran family Zhangsolvidae. This speculation needs additional corroboration from other groups, that will become available with the combination of systematics, paleontology and biogeographical information of different early Cretaceous clades.     

New snakeflies (Insecta: Raphidioptera) from the Lower Cretaceous of the UK,Spain and Brazil

Abstract: Several new taxa of snakeflies (Raphidioptera) are described from the Lower Cretaceous deposits of the Wealden, UK (Barremian), Montsec, Spain (Barremian) and the Crato Formation, Brazil (Aptian). Mesoraphidia ednae sp. nov. and M. hilli sp. nov. are described from the Wealden; Nanoraphidia lithographica sp. nov. and Iberoraphidia dividua gen. et sp. nov. are described from Montsec, and Baissoptera lisae sp. nov. is described from the Crato Formation. The geographical range of Nanoraphidia has potentially been extended.     

Cratonia cotyledon gen. et sp. nov: a unique Cretaceous seedling related to Welwitschia

The fossil history of most extant seed plant groups is relatively well documented. Cycads, conifers and Ginkgo all have an extensive fossil record, and the understanding of early angiosperm diversity is increasing. The Gnetales are an exception. Few macrofossils have been described, and character evolution within the group is poorly known. Cratonia cotyledon is a new gnetalean fossil from the Early Cretaceous Crato Formation of Brazil. This well-preserved seedling consists of two cotyledons, a feeder and a root. The leaf surface shows polygonal epidermal cells and apparently paracytic or actinocytic stomata. The cotyledons have a very specific venation pattern, shared only by Cratonia and Welwitschia, with parallel primary veins and secondary veins fusing to form inverted 'Y's between the main veins. Based on the 'Y'-venation and the presence of a feeder, we assign Cratonia to the Gnetum-Welwitschia clade. Fossil seedlings are unusual and this complete specimen with unambiguously welwitschioid characters is spectacular. Cratonia indicates that the evolutionary split between Gnetum and Welwitschia had occurred in the Early Cretaceous. Further, the close relationship between a West African plant and an east South American Early Cretaceous fossil is consistent with a major geological event: the rifting of the Gondwana continent.     

New freshwater mussels (Bivalvia,Unionida) with potential trigonioidid and hyriid affinities from the Early Cretaceous of Brazil

Two new taxa of freshwater mussels (Bivalvia: Unionida) from the Aptian Crato Formation of the Araripe Basin, NE Brazil, are described. The fossil bivalves are confined to 30- to 130-cm-thick bioturbated mudstones overlying the fossil-rich laminated limestones of the Crato Formation Konservat-Lagerstätte. Individuals are often preserved with closed or splayed articulated valves, some of them potentially in life position, forming an autochthonous to parautochthonous assemblage. Monginellopsis bellaradiata nov. gen., nov. sp. shares key characters with the Trigonioidoidea: (i) the anterior pedal retractor muscle scar is clearly separated from the anterior adductor muscle scar; (ii) the shell has fold-like radial ribs on the posterior half; (iii) a right valve anterior tooth has a striated facet. Araripenaia elliptica nov. gen., nov. sp. is the most abundant and widely distributed unionid of the Crato Formation. Its ornament of anterior inverted V-shaped riblets, and central and posterior radial and sub-radial riblets resembles modern and fossil Hyriidae from the Americas, but also Trigonioidoidea from Eurasia. Its dentition of two smooth anterior pseudocardinals and two smooth posterior laterals in each valve provides no further clues for systematic assignment; muscle scars are not preserved. Assignment to the Hyriidae would make Araripenaia the oldest member of this family known from South America. Moreover, this bivalve assemblage of trigonioidoidids, hyriids, and previously reported silesunionoids suggests palaeobiogeographic links to other areas in both Gondwana and Laurasia.     

Small Celastraceae and Polygonaceae twigs from the Upper Cenozoic (Ituzaingó Formation) of the La Plata Basin,Argentina

Two new wood types from the Late Cenozoic of the Ituzaingó Formation, La Plata Basin, Northeast Argentina add to our knowledge of South American Cenozoic plants. The materials were preserved by siliceous cellular permineralization, and they were prepared for microscopic examination by surface polishing and in thin sections. The anatomy of these new species was described. The relationship and comparison with the nearest living relatives (NLRs) are discussed. Maytenoxylon perforatum Franco gen. and sp. nov. is described as the first fossil wood referable to Celastraceae from South America. This new fossil species is related to extant Maytenus Molina. The other fossil twig, Ruprechtioxylon breae Franco sp. nov., has features of the Polygonaceae family and particularly resembles the extant specie Ruprechtia laxiflora Meisn. The occurrence of these fossil woods in south-eastern South America suggests that a relatively warm and dry to seasonally dry climate prevailed over this region of Gondwana during the Upper Cenozoic. It also provides new evidence for the hypothesis of the more wide distribution of Seasonally Dry Tropical Forest (SDTF) during the Upper Cenozoic.     

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 genus and species of carrion beetle (Coleoptera,Silphidae) from the Lower Cretaceous Jinju Formation,South Korea

A new fossil genus of Silphidae, Cretosaja gen. nov., is described, based on a new fossil species, Cretosaja jinjuensis sp. nov. from the Lower Cretaceous Jinju Formation in South Korea. The association of Cretosaja with Nicrophorinae is discussed on the basis of morphological evidence. Diagnostic differences of the fossil and extant genera of Nicrophorinae are provided. Primitive characteristics presented in Mesozoic silphids are discussed.     

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     

THE FIRST FOSSIL PROSCOPIIDAE (INSECTA, ORTHOPTERA, EUMASTACOIDEA) WITH COMMENTS ON THE HISTORICAL BIOGEOGRAPHY AND EVOLUTION OF THE FAMILY

Abstract:  Eoproscopia martilli gen. et sp. nov. is described from the Early Cretaceous (Aptian) Crato Formation Lagerstätte of Ceará State, north-east Brazil. The new taxon is assigned to the extant family Proscopiidae and represents the first occurrence of the group in the fossil record. Eoproscopia is similar to crown group proscopiids in its stick-like habitus, elongate prothorax and absence of the cryptopleuron, but differs in the presence of well-developed wings, the short head with a small, simple fastigium, the prothoracic legs being inserted near the posterior margin of the prothorax, and the absence of spines on the metathoracic tibiae. The discovery of Eoproscopia extends the geological range of the family by approximately 110 myr and confirms the presence of stem-group proscopiids in the Atlantic rift zone of South America during the Early Cretaceous.     

A New Japygid Dipluran from the Lower Cretaceous of Brazil

The first Mesozoic japygid (Hexapoda: Diplura), Ferrojapyx vivax gen. et sp. nov., is described from the Lower Cretaceous (Aptian) Crato Formation of north-east Brazil. There are only two previously described occurrences of japygids in the fossil record: in Miocene or Pliocene onyx marble from Arizona and from the Carboniferous (Westphalian D) Francis Creek Shale of Mazon Creek, Illinois.     

A specimen of Curculioninae (Curculionidae,Coleoptera) from the Lower Cretaceous,Araripe Basin,north‐eastern Brazil

Abstract: A specimen of Curculionidae (Curculioninae) is described as Arariperhinus monnei gen. et sp. nov. The specimen is preserved on a laminated limestone sample of the Crato Formation (Santana Group), Lower Cretaceous (Aptian–Albian), and was collected from a quarry near Nova Olinda, Chapada do Araripe, State of Ceará, Brazil. The genus is placed in the subfamily Curculioninae because of its strongly convex body and relatively slender rostrum, but mainly by its rounded eyes and lack of a prosternal sulcus and tibial spurs. The very prominent eyes in lateral view, a cylindrical rostrum and a straight posterior margin of ventrite II are strong indications that this fossil belongs to the tribe Anthonomini. However, the claws, which would resolve the exact placement of this fossil, are poorly preserved. Arariperhinus monnei gen. et sp. nov. is distinguishable by the combination of several characters and the first record of the family Curculionidae in the Santana Group; it is the oldest record of a member of the subfamily Curculioninae.     

First Mesozoic Scutigeromorph Centipede, from the Lower Cretaceous of Brazil

The first Mesozoic scutigeromorph centipede (Myriapoda: Chilopoda), Fulmenocursor tenax gen. et sp. nov., is described from the Lower Cretaceous (Aptian) Crato Formation of north-east Brazil. Previously described fossil Scutigeromorpha are known from Dominican and Baltic amber, the Carboniferous (Westphalian D) Francis Creek Shale of Mazon Creek, Illinois, the Silurian and Devonian of Britain, and the Devonian of New York State.     

NEW FOSSIL LACEWINGS AND ANTLIONS (INSECTA, NEUROPTERA) FROM THE LOWER CRETACEOUS CRATO FORMATION OF BRAZIL

Abstract:  Remarkable new fossil taxa of Neuroptera from the laminated limestone of the Crato Formation, north-east Brazil, are described: Nuddsia longiantennata gen. et sp. nov. (Osmylidae, Gumillinae), the first fossil record of this family from South America, Parapalaeoleon magnus gen. et sp. nov. (Palaeoleontidae), and Triangulochrysopa formosa sp. nov. (Mesochrysopidae). A diagnosis of Gumillinae is provided; Epiosmylidae is considered to be a synonym of this subfamily. A revised diagnosis of Palaeoleontidae is provided, based on a new interpretation of venational characters evident on the exceptionally well-preserved specimen of P. magnus ; vein homologies are determined and the fusion of MP and CuA is discussed. The genus Triangulochrysopa has been known previously only from the Lower Cretaceous of Las Hoyas, Spain.     

A new genus of Staphylinidae (Coleoptera) from the Lower Cretaceous: the first fossil rove beetles from the Southern Hemisphere

The fossil genus of rove beetles Apticax gen.n . with two new species, A. volans sp.n . and A. solidus sp.n., is described from the Nova Olinda Member of the Crato Formation in north‐eastern Brazil (Aptian–Albian, dated as 125–99.6 Ma old). Both species belong to the clade Staphylininae + Paederinae in the staphylinine group of subfamilies and are the first fossil true Staphylinidae to be described from the entire Southern Hemisphere. Although they resemble Paederinae, the low number of satisfactorily preserved characters do not allow definite placement of either A. volans or A. solidus in any of the subfamilies of Staphylinidae.     

A NEW AZHDARCHOID PTEROSAUR FROM THE CRATO FORMATION (LOWER CRETACEOUS,APTIAN?) OF BRAZIL

Abstract: A partial pterosaur skull from the Nova Olinda Member of the Crato Formation (Lower Cretaceous, Aptian?) represents a new edentulous pterodactyloid, Lacusovagus magnificens gen. et sp. nov. The absence of teeth and a large nasoantorbital fenestra suggest assignment to Azhdarchoidea, and the combination of a particularly short, crestless and shallow rostrum and laterally flared jaw margins distinguish it from other azhdarchoid taxa. The position of the new form within Azhdarchoidea is problematic: Lacusovagus is distinguished from Tapejaridae in its straight, as opposed to ventrally displaced, jaw tip and absence of a premaxillary crest; from thalassodromids by the absence of a premaxillary crest; and from Azhdarchidae by the short length of the rostrum and shallow posterodorsal extension of the premaxilla. Lacusovagus shares a shallow, crestless rostrum and a slender posterodorsal premaxillary extension with Jiufotang Formation azhdarchoids such as Chaoyangopterus and Jidapterus. The position of these genera within Azhdarchoidea is controversial, but the suite of plesiomorphic and derived azhdarchoid characters in each suggests a placement between Tapejaridae and Neoazhdarchia. Further research is required, however, to determine the relationships of these genera both to each other and to other azhdarchoids. The new taxon elevates the faunal similarity found between the roughly contemporaneous Jiufotang and Crato formations and continues the pattern of Crato Formation azhdarchoids being much larger than those from the Jehol Group. It also has jaws at least 67 and 55 per cent longer, respectively, than those of the largest azhdarchoids and ornithocheirids from the Crato pterosaur assemblage, making Lacusovagus the largest pterosaur known from this unit.     

The oldest species of the genus <Emphasis Type="Italic">Glaphyrus</Emphasis> Latr. (Coleoptera: Scarabaeoidea: Glaphyridae) from the Mesozoic of China

Glaphyrus ancestralis sp. nov. is described from the Yixian Formation (Upper Jurassic or Lower Cretaceous). The species is not only one of the earliest records of the family Glaphyridae but also the oldest representative of an extant genus of the family.     

Marsileaceaephyllum, a new genus for marsileaceous macrofossils: leaf remains from the Early Cretaceous (Albian) of southern Gondwana

Three new taxa from Albian, Early Cretaceous assemblages in Gondwana (Australia and Antarctica) and two previously described fossils from the Late Cretaceous and Eocene of North America are attributable to the heterosporous semi-aquatic fern family Marsileaceae. They are assigned to Marsileaceaephyllum, a morphotaxon erected here for sterile remains (whole plants, and isolated leaves and leaflets) of Marsileaceae. The Gondwanan taxa, Marsileaceaephyllum lobatum and Marsileaceaephyllum spp. B-C, have either a cruciform leaflet arrangement or dichotomous and anastomosing venation characteristic of modern Marsileaceae. Two previously established taxa, Marsilea johnhallii and Marsilea sp., which represent sterile Marsileaceae, are also transferred to the new genus (now Marsileaceaephyllum johnhallii and Marsileaceaephyllum sp. A, respectively). Examination of all fossil venation patterns reveals four new venation types not present in extant taxa, suggesting that most fossil Marsileaceae (leaves) are distinct from extant genera, and are likely members of extinct lineages. This is further supported by the absence of modern megaspore types in the Early Cretaceous.     

First fossil record of the genus Nageia (Podocarpaceae) in south China and its phytogeographic implications

The first fossil record of the genus Nageia Gaertner from the Tertiary of southern China is reported. Nageia hainanensis sp. nov. is described from the Eocene Changchang Formation of Hainan Island. The extant species of Nageia are widely distributed in southeastern Asia, from northeastern India to southeastern China and southern Japan, and southward through Indochina to the Malay Archipelago and New Guinea. The fossil evidence of Nageia has been discovered from the Lower Cretaceous of the Far East of Russia and Japan. The distribution of both modern and fossil species of Nageia indicates that this genus could have originated in the northeastern part of Asia in the Early Cretaceous and spread to south China at least in the Eocene. Discovery of Nageia fossil record in south China provides important evidence for the geological and phytogeographic history of the genus.     

HIGHER SYSTEMATICS OF SCORPIONS FROM THE CRATO FORMATION, LOWER CRETACEOUS OF BRAZIL

Abstract:  Several new specimens of Protoischnurus axelrodorum Carvalho and Lourenço and Araripescorpius ligabuei Campos from the Crato Formation, Brazil, are described. The preservation and recognition of new morphological features allows a re-diagnosis of both species and a modification of their familial placement. Protoischnurus axelrodorum is the oldest species belonging to the scorpionoid family Hemiscorpiidae Pocock (= Ischnuridae Simon; = Liochelidae Fet and Bechly) and the first Cretaceous record. It was originally placed in the extinct family Protoischnuridae Carvalho and Lourenço, which is here synonymized with Hemiscorpiidae. Araripescorpius ligabuei, now assigned to Chactidae Pocock, is the first chactoid recorded for the Cretaceous of Brazil. These findings confirm that the lineages of two modern families date back at least to the Early Cretaceous and, considering their current distribution, were probably present before the break-up of the supercontinent Gondwana. Palaeoecological inferences indicate the presence of tropical habitats in the vicinity of the Crato lake/lagoon.     

A scolopocryptopid centipede (Chilopoda: Scolopendromorpha) from Mexican amber: synchrotron microtomography and phylogenetic placement using a combined morphological and molecular data set

The first scolopocryptopid centipede known from the fossil record is a specimen of the subfamily Scolopocryptopinae in Miocene amber from Chiapas, southern Mexico. It is described here as Scolopocryptops simojovelensis sp. nov. , displaying a distinct combination of morphological characters compared to extant congeners. Anatomical details of the fossil specimen were acquired by non‐invasive 3D synchrotron microtomography using X‐ray phase contrast. The phylogenetic position of the new species is inferred based on a combination of morphological data with sequences for six genes (nuclear 18S and 28S rRNA, nuclear protein‐coding histone H3, and mitochondrial 12S rRNA, 16S rRNA, and protein‐coding cytochrome c oxidase subunit I) for extant Scolopendromorpha. The data set includes eight extant species of Scolopocryptops and Dinocryptops from North America, east Asia, and the Pacific, rooted with novel sequence data for other blind scolopendromorphs. The molecular and combined data sets, analysed in a parsimony/direct optimization framework, identified a stable pattern of two main clades within Scolopocryptopinae. North American and Asian species of Scolopocryptops are united as a clade supported by both morphological and molecular characters. Its sister group is a Neotropical clade in which the type species of Dinocryptops is nested within a paraphyletic assemblage of Scolopocryptops species; Dinocryptops is placed in synonymy with Scolopocryptops. The strength of support for the relationships of extant taxa from the molecular data allow the Chiapas fossil to be assigned with precision, despite ambiguity in the morphological data; the fossil is resolved as sister species to the extant Laurasian clade. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 768–786.