French Amber Research Articles

Barklice (Insecta: Psocodea) from Early Cretaceous resiniferous forests of Iberia (Spanish amber): New Troctomorpha and a possible Psocomorpha

Psocids, commonly known as barklice, are insects belonging to the order Psocodea, together with the parasitic lice. They usually inhabit forest litter or the bark of tree trunks and branches, showing grazing herbivorous or detritivorous feeding habits. The Cretaceous psocid record is diverse, containing more than 70 described species. Here, we present new psocids (Troctomorpha and possible Psocomorpha) from two Spanish amber outcrops, both Albian (Lower Cretaceous): El Soplao and Ariño. We describe the two new species Azarpsocus anjana Álvarez-Parra and Nel sp. nov. (Manicapsocidae) and Burmacompsocus ojancano Álvarez-Parra and Nel sp. nov. (Compsocidae), and the morphotype Ariño that might belong to Psocomorpha. We discuss the taxonomic placement of the studied specimens and comment on the palaeobiogeography of the Cretaceous psocids. The Cretaceous barklice fauna from Iberia shows more similarities with that from Lebanese (Barremian) and Burmese (Cenomanian) ambers than with that from the palaeogeographically closer French amber (Cenomanian). This finding has been also reported in other insect groups from Spanish amber, and might be the consequence of several factors, such as the palaeoenvironmental conditions of the ecosystems, the palaeogeographical position of the Iberia Island during the latest Jurassic, closer to Gondwana than to Laurasia, and oceanic currents that facilitated or hindered the displacement of insect fauna between continental masses.

Cretaceous Antodicranomyia (Diptera: Limoniidae) and their paleohabitat

New representatives of the Cretaceous cranefly genus Antodicranomyia (Diptera: Limoniidae) are reported from Albian-Cenomanian Charentese (French) amber. The newly reported specimens allow for an emended diagnosis of the type species A. azari, as well as the description of a new species, Antodicranomyia rubra sp. nov., which is mostly distinguished from the type species by features of its wing venation, antennae, and genitalia. As a rare, extinct genus known only from French amber, Antodicranomyia is compared with its closest relative genera Antocha, Dicranomyia and Antohelia. The evolutionary implications and paleohabitat of Antodicranomyia are discussed. The new discovery adds to the knowledge of the crane flies’ diversity and evolution in the mid-Cretaceous.

A previously missing link in the evolution of dasytine soft‐winged flower beetles from Cretaceous Charentese amber (Coleoptera, Melyridae)

AbstractThe soft‐winged flower beetles are a diverse cosmopolitan family of cleroid beetles with an unresolved higher classification, the adults of which feed on nectar and pollen. The fossil record of Melyridae is sparse, with only two described fossils known from the Mesozoic. Here we describe the first representative of the family from the Mesozoic of Europe, from early Cenomanian Charentese (French) amber. Due to the opaque nature of French amber, X‐ray microtomography was used to document fine morphological details of the fossil. Protodasytes cretaceus gen. et sp. nov. possesses a unique combination of characters within the subfamily Dasytinae, which is shared by the tribes Chaetomalachiini, Dasytini, and Listrini, namely the presence of a small antennal club, tarsomere 4 slightly shorter than 3 but not distinctly narrower, symmetrical claws, and a distinct pronotal and elytral fringe. This suggests that Dasytinae had begun to diversify by the Late Cretaceous, possibly in concert with the radiation of angiosperms. The fossil record of soft‐winged flower beetles is critically reviewed, and a checklist of extinct species is provided. The fragmentary Cenozoic fossils Troglops punctatissima and Malachius vertummni are removed from the family Melyridae, and the record of Attalus lusitanicus from Eocene–Oligocene Bembridge Marls is regarded as unlikely. The potential role of Protodasytes gen. nov. and Cretaceous melyrids as pollinators of early angiosperms is discussed.

Ankothrips, the most ancient extant thrips genus (Thysanoptera, Melanthripidae)

AbstractTwo fossil representatives of the extant genus Ankothrips are described from French amber, namely, the Late Cretaceous Ankothrips dupeae sp. nov., from Vendean amber, and the earliest Eocene Ankothrips deploegi sp. nov., from Oise amber. We discuss the differences between the middle Eocene Baltic amber genus Archankothrips and Ankothrips, concluding that the two genera are extremely similar with only one significant difference in male specimens. The disjunctive distribution of the extant species along with the presence of one Late Cretaceous species strongly suggest an Early Cretaceous origin for Ankothrips. The genus is therefore very ancient and has had a remarkable morphological stability for the last 94 million years. This situation is unique among the Cretaceous thrips, and rare among insects (only c. 3% of the genera in the well‐documented mid‐Cretaceous Burmese amber are still extant). We propose some hypotheses that could explain the stability of Ankothrips through time.

100-million-year-old ant–conifer associates inside French amber: a fortuitous or ecological association?

100-million-year-old ant–conifer associates inside French amber: a fortuitous or ecological association?

The oldest representative of the bombyliid bee fly in the earliest Eocene French amber

The oldest representative of the bombyliid bee fly in the earliest Eocene French amber

A new species ofAmpulicomorphaAshmead from Eocene French amber, with a list of fossil and extant Embolemidae (Insecta: Hymenoptera) of the world

Embolemidae is a cosmopolitan but species-poor group of chrysidoid wasps with a scarce fossil record, despite a long evolutionary history since at least the Early Cretaceous. Here, the new species,Ampulicomorpha quesnoyensissp. nov., is illustrated and described based on a single female found in Early Eocene amber of Oise (France). The new species is compared with the three other known fossil species of the genus, and a key to all fossil species ofAmpulicomorphais provided. This is the third European fossil species ofAmpulicomorpha, which suggests that the genus was once well established in Western Europe while it is more widely distributed in the Eastern Palaearctic region today. A list of all fossil and extant Embolemidae of the world, as well as a map of their geographical distribution map, are provided.

A new species of the haematophagous genus Austroconops Wirth & Lee (Diptera: Ceratopogonidae: Leptoconopinae) from middle Cretaceous amber of Charente-Maritime, France

Austroconops perrichoti, a new species of biting midge (Diptera: Ceratopogonidae), is described from uppermost Albian-lowermost Cenomanian amber of Charente-Maritime (NW France), from three female specimens. A similar species, Austroconops borkenti Szadziewski & Schlüter, 1992, from Upper Cretaceous, Cenomanian, French amber of Anjou, is redescribed and illustrated. A key to males and females of extinct species of Austroconops Wirth & Lee is also provided.

The semi-aquatic pondweed bugs of a Cretaceous swamp.

Pondweed bugs (Hemiptera: Mesoveliidae), considered a sister group to all other Gerromorpha, are exceedingly rare as fossils. Therefore, each new discovery of a fossil mesoveliid is of high interest, giving new insight into their early evolutionary history and diversity and enabling the testing of their proposed relationships. Here, we report the discovery of new mesoveliid material from Spanish Lower Cretaceous (Albian) amber, which is the first such find in Spanish amber. To date, fossil records of this family only include one species from French Kimmeridgian as compression fossils, two species in French amber (Albian-Cenomanian boundary), and one in Dominican amber (Miocene). The discovery of two males and one female described and figured as Glaesivelia pulcherrima Sánchez-García & Solórzano Kraemer gen. et sp. n., and a single female described and figured as Iberovelia quisquilia Sánchez-García & Nel, gen. et sp. n., reveals novel combinations of traits related to some genera currently in the subfamily Mesoveliinae. Brief comments about challenges facing the study of fossil mesoveliids are provided, showing the necessity for a revision of the existing phylogenetic hypotheses. Some of the specimens were studied using infrared microscopy, a promising alternative to the systematic study of organisms preserved in amber that cannot be clearly visualised. The new taxa significantly expand the fossil record of the family and shed new light on its palaeoecology. The fossils indicate that Mesoveliidae were certainly diverse by the Cretaceous and that numerous tiny cryptic species living in humid terrestrial to marginal aquatic habitats remain to be discovered. Furthermore, the finding of several specimens as syninclusions suggests aggregative behaviour, thereby representing the earliest documented evidence of such ethology.

Palaeobiology of tanaidaceans (Crustacea: Peracarida) from Cretaceous ambers: extending the scarce fossil record of a diverse peracarid group

Diverse assemblages of tanaidacean peracarid crustaceans from western Tethyan continental deposits suggest that the group was relatively common in or around ancient resin-producing forests. Here we report the results of an examination of 13 tanaidacean specimens from three Cretaceous (Albian to Turonian) French amber deposits. Two new species of the fossil family Alavatanaidae are placed in the previously described Early Cretaceous genus Eurotanais: Eurotanais pyrenaensis sp. nov. from Cenomanian Pyrenean amber (Fourtou, Aude) and Eurotanais seilacheri sp. nov. from Turonian Vendean amber (La Garnache, Vendee). The remaining specimens are placed in three newly erected genera and species (but family incertae sedis): Arcantitanais turpis gen. et sp. nov. from Albian–Cenomanian Charentese amber (Archingeay, Charente-Maritime), and Tytthotanais tenvis gen. et sp. nov. and Armadillopsis rara gen. et sp. nov. from Pyrenean amber. These are the first formally described fossils that might be related to the paratanaoidean families Nototanaidae and Paratanaidae, sharing with these some putatively derived features and providing possible evidence for the antiquity and morphological stability of these families and the suborder Tanaidomorpha. The distinctive features and character combinations of these fossil taxa are discussed in connection with possible relationships to the living lineages of tanaidaceans. Propagation phase-contrast X-ray synchrotron microtomography was used to obtain high-quality 3D images for some fossils. A discussion is provided on the putative palaeobiology of tanaidaceans and the French resiniferous forest ecosystem. The discovery of these new tanaidaceans extends the palaeogeographical distribution and stratigraphical range of the family Alavatanaidae and sheds new light on the palaeoecology and diversity of tanaidaceans in pre-angiospermous woodlands.

Evolutionary and paleobiological implications of Coleoptera (Insecta) from Tethyan-influenced Cretaceous ambers

The intense study of coleopteran inclusions from Spanish (Albian in age) and French (Albian–Santonian in age) Cretaceous ambers, both of Laurasian origin, has revealed that the majority of samples belong to the Polyphaga suborder and, in contrast to the case of the compression fossils, only one family of Archostemata, one of Adephaga, and no Myxophaga suborders are represented. A total of 30 families from Spain and 16 families from France have been identified (with almost twice bioinclusions identified in Spain than in France); 13 of these families have their most ancient representatives within these ambers. A similar study had previously only been performed on Lebanese ambers (Barremian in age and Gondwanan in origin), recording 36 coleopteran families. Few lists of taxa were available for Myanmar (Burmese) amber (early Cenomanian in age and Laurasian in origin). Coleopteran families found in Cretaceous ambers share with their modern relatives mainly saproxylic and detritivorous habits in the larval or adult stages, rather than wood-boring behavior. Fifteen of the coleopteran families occur in both the Lebanese and Spanish ambers; while only five are present in both Spanish and French. Considering the paleogeographic proximity and similarity of age of the Spanish and French ambers, the small number of taxa found in common at both areas is surprising. The ancient origin for the Lebanese and Spanish ambers, the paleogeography (including some barriers for terrestrial biota) and the local paleohabitats are factors that may explain the dissimilarity with the French specimens. Wildfires are believed to be a more likely cause of resin production during the Cretaceous than infestation by beetles. Current knowledge of the beetle species found in the Cretaceous ambers is introduced.

A new species of the genus Clidicus from Lower Cretaceous of France (Coleoptera: Staphylinidae: Scydmaeninae)

Clidicus cretaceus sp. nov. from the Lower Cretaceous French amber of Archingeay is described. This finding is important to correct the phylogenetic hypotheses proposed for the tribe Clidicini.

Biting midges of the extinct genus Protoculicoides Boesel from Lower Cretaceous amber of San Just, Spain and new synonymy in recently described fossil genera (Diptera: Ceratopogonidae)

We describe two new species of extinct biting midges in the fossil genus Protoculicoides: Protoculicoides hispanicus Szadziewski and Arillo, sp. nov. and Protoculicodes sanjusti Szadziewski and Arillo, sp. nov., from Albian (101–113 Ma) amber of San Just, Spain. Atriculicoides Remm, 1976 is recognized as a new junior synonym of Protoculicoides Boesel, 1937 (NEW SYNONYM). Devalquia Choufani et al., 2013 is regarded here as a new junior synonym of the extant genus Culicoides Latreille, 1809 (NEW SYNONYM). Metahelea roggeroi Choufani et al., 2013 from Upper Cretaceous French amber, is excluded from the tribe Heteromyiini and placed in the extant genus Stilobezzia Kieffer, 1911, tribe Ceratopogonini: Stilobezzia roggeroi (Choufani et al., 2013), comb. nov. The extinct tribe Atriculicoidini Szadziewski, 1996 is regarded as a new subfamily, Atriculicoidinae, stat. nov., that includes the fossil genus Protoculicoides, with 13 species reported from Albian, Turonian, Cenomanian, Coniacian, Santonian and Campanian ambers (78–113 Ma). Protoculicoides skalskii Szadziewski & Arillo, 1998 from Lower Cretaceous amber of Álava, Spain, Protoculicoides succineus Szadziewski, 1966 from Lower Cretaceous Lebanese amber and Protoculicoides burmiticus Szadziewski & Poinar, 2005 from Upper Cretaceous Burmese amber are transferred to the extinct genus Archiculicoides Szadziewski, 1996, comb. nov. Protoculicoides krzeminskii Choufani et al., 2014 from Lower Cretaceous Lebanese amber is transferred to the extinct genus Archiaustroconops Szadziewski, 1996, comb. nov. The Cretaceous subfamily Atriculicoidinae forms an unresolved trichotomy with the extant subfamilies Forcipomyiinae and Dasyheleinae, both of which date to the Eocene. The fossil record indicates that wings with macrotrichia in biting midges evolved during the mid-Cretaceous greenhouse climate. We suggest that this was an evolutionary adaptation to new atmospheric conditions with higher levels of CO2 in order for the antennal Johnston's organs of males to receive the vibrational sex signals produced by females during flight.

Chemical Characterization and Botanical Origin of French Ambers.

The molecular composition of 10 Cretaceous and one Eocene ambers from France was analyzed by infrared spectroscopy, solid-state (13)C nuclear magnetic resonance spectroscopy, and thermochemolysis gas chromatography-mass spectrometry. The terpenoids identified in the samples were used as biomarkers for the botanical origin of the resins. The Cretaceous samples, comprising the so-called Alpine, Anjou, Charentese, Provence, Pyrenean, and Vendean ambers, ranged from the Albian-Cenomanian transition to the early Santonian (100 to 85 Ma) and correspond to class Ib resins typical of conifers. The extinct conifer family Cheirolepidiaceae was proposed as the plant source of Pyrenean and brown Charentese ambers. Araucariaceae or Cheirolepidiaceae were the plant sources of the Cenomanian Alpine, Anjou, and yellow Charentese ambers. The Santonian ambers of Provence and Vendée were found to derive from the Cupressaceae. The Eocene Oise amber (ca. 53 Ma) is a class Ic resin typical of angiosperms and was produced by a Fabaceae. The evolution of resin sources from the early Cretaceous to the Eocene periods is discussed. Finally, a possible fingerprint hitherto unveiled is proposed for cheirolepidiaceous resins, defined by the simultaneous presence of phenolic diterpenoids, labdanoic acids, callitrisate structures, and their respective derivatives.

Spider web in Late Cretaceous French amber (Vendée): The contribution of 3D image microscopy

Late Cretaceous amber from La Garnache (France, Vendée) contains filamentous networks that have all the characteristics of spider webs. Using methods of classic and confocal microscopy, the web architecture and the thread structure are described. The geometry of the webs preserved in amber is presented in 3D imaging. Unlike most of the spider webs identified in amber, there are no regular radiating webs growing in two dimensions characteristic of the well-known orb web. A number of the characters would correspond to the apparently irregular organization of cob-type web or sheet web. By using confocal microscopy, we can highlight the preserved autofluorescence of spider silk proteins and new characters are proposed to identify the spider webs or fragments of webs in amber.

A new trap-jawed ant (Hymenoptera: Formicidae: Haidomyrmecini) from Canadian Late Cretaceous amber

AbstractA new genus and species are described within the extinct tribe Haidomyrmecini, and tentatively placed within the subfamily Sphecomyrminae (Hymenoptera: Formicidae).Haidoterminus cippusnew genus and speciesexpands the distribution of the bizarre, exclusively Cretaceous, trap-jawed Haidomyrmecini beyond their previous records in mid-Cretaceous Burmese and French amber, and into Laurentia. The new material from the Grassy Lake, Alberta, Canada collecting locality also provides evidence that these highly specialised, likely arboreal, ants persisted for an additional 20 million years, reaching the Late Cretaceous. Morphological features ofH. cippus, such as the presence of an elongate antennomere II (pedicel), further support the argument that Haidomyrmecini may not actually belong within the subfamily Sphecomyrminae, and may warrant recognition at the subfamily level or inclusion as a highly autapomorphic clade within another subfamily. Despite the introduction of new fossil material, and the clarity of preservation in Canadian amber, the mystery of how Haidomyrmecini fed remains unsolved.

New maimetshid wasps in Cretaceous amber from Myanmar (Insecta: Hymenoptera)

The extinct, exclusively Cretaceous wasp family Maimetshidae is newly recorded from Earliest Cenomanian Burmese amber. Two new genera and species are described. Burmaimetsha concava gen. et sp. nov., based on a male and a female, is most similar to Guyotemaimetsha Perrichot, Nel & Néraudeau, from Albian-Cenomanian French amber, but differs in its larger mandibles, distinctly concave face, elongate antennomeres, and forewing with cell [1Rs] smaller and fourth abscissa of Rs shorter. Maimetshasia kachinensis gen. et sp. nov., is based on a male, and is characterized by asymmetric mandibles with two and three teeth, by its forewing venation without cross-vein 2rs-m, with cell [1M] large and trapezoidal, and vein 2Rs+M very short, and by the hind wing without a free apex of Rs. The family was evidently widespread in the Cretaceous, and the new records extend the paleobiogeographical range to the South-East of Eurasia. A discussion about the possible biology of Maimetshidae is provided.

Rediscovery of the Bizarre Cretaceous AntHaidomyrmexDlussky (Hymenoptera: Formicidae), with Two New Species

ABSTRACT The discovery of two distinct, near-complete specimens belonging to the Cretaceous ant genus Haidomyrmex Dlussky prompts a detailed description and discussion of a remarkable mandibular morphology. The specimens, preserved in 98 million-year-old amber from northern Myanmar, are described here as Haidomyrmex scimitarus, n. sp., and Haidomyrmex zigrasi, n. sp., with diagnostic differences provided between them as well as with H. cerberus Dlussky (also in Burmese amber). Relationships and comparisons of H. scimitarus, H. zigrasi, H. cerberus, and the recently described Haidomyrmodes mammuthus Perrichot from Cretaceous French amber are also discussed. Haidomyrmex was probably arboreal, cursorial, and a specialized trap-jaw predator, utilizing its enormous mandibles and cranial morphology in concert to capture prey. Mandibles appear to have moved in a plane oblique to the dorsoventral and horizontal axes of the body, unlike the lateral-plane movement of modern ants. The additions of these new fossils ...

Fossil amoebae (Hemiarcherellidae fam. nov.) from Albian (Cretaceous) amber of France

Abstract: Two extraordinarily well‐preserved testate amoebae are described from Late Albian age amber from south‐western France. The specimens are attributed to a new family, the Hemiarcherellidae fam. nov., and are described as Hemiarcherella christellae gen. et sp. nov. The amoebae described herein originate from highly fossiliferous amber pieces. Based on syninclusions, Hemiarcherella christellae was a soil‐dwelling organism, probably an active bacterivore. This taxon represents the third species of testate amoebae described from mid‐Cretaceous French amber. Analysis of this fossil amoeba fauna illustrates the uniqueness of mid‐Cretaceous French amber deposits. Indeed, most amoebae found in amber have been assigned to modern species, corroborating the hypothesis of morphological stasis in different microbial lineages. However, the well‐preserved amoebae fauna found in French amber can be distinguished clearly from modern species and help us to better understand the fossil record of these organisms.

The French ambers: a general conspectus and the Lowermost Eocene amber deposit of Le Quesnoy in the Paris Basin

Aconspectus of the current knowledge on the French ambers is proposed. France is a very rich country with more than seventy localities known after the 'old' literature, especially from the Jurassic, Cretaceous and Paleogene. A more detailed new insectiferous amber locality is given for the amber from Le Quesnoy (Lowermost Eocene, Oise department, Paris Basin, France). After the preliminary survey of the flora and the vertebrate and arthropod faunas, we propose a reconstruction of a fluvio-lacustrine palaeoenvironment with a forest, under a warm and wet seasonal climate. This site is outstanding because of the richness, diversity and the state of preservation of the fossils. The present discovery opens a remarkable window on the terrestrial life during the earliest Eocene. It shows that future researches on the 'old' French amber localities shall be crucial for our knowledge on the evolution and diversity of the insects during the Mesozoic and Early Cenozoic.