Xavier Delclòs

Thrips pollination of Mesozoic gymnosperms

Within modern gymnosperms, conifers and Ginkgo are exclusively wind pollinated whereas many gnetaleans and cycads are insect pollinated. For cycads, thrips are specialized pollinators. We report such a specialized pollination mode from Early Cretaceous amber of Spain, wherein four female thrips representing a genus and two species in the family Melanthripidae were covered by abundant Cycadopites pollen grains. These females bear unique ring setae interpreted as specialized structures for pollen grain collection, functionally equivalent to the hook-tipped sensilla and plumose setae on the bodies of bees. The most parsimonious explanation for this structure is parental food provisioning for larvae, indicating subsociality. This association provides direct evidence of specialized collection and transportation of pollen grains and likely gymnosperm pollination by 110–105 million years ago, possibly considerably earlier.

Mesozoic chrysopid-like Planipennia: a phylogenetic approach (Insecta: Neuroptera)

Abstract The Mesozoic chrysopid-like Planipennia are revised and several new genera and species are described. The new superfamily Chrysopoidea is proposed for the extant and fossil Chrysopidae, and the fossil families Liassochrysidae n. fam., Allopteridae Zhang 1991 n. sensu, Mesochrysopidae Handlirsch, 1906 n. sensu, Tachinymphidae n. fam., and Limaiidae Martins-Neto and Vulcano 1989 n. sensu. A phylogenetic analysis of the Chrysopoidea is proposed, based on the wing venation characters. With at least the four families Allopteridae, Mesochrysopidae, Tachinymphidae, and Chrysopidae, showing different wing venation patterns, the systematic diversity and morphological disparity of the Chrysopoidea are maximal during the Late Jurassic and Early Cretaceous. The Mesozoic family Limaiidae was still present during the Paleocene/Eocene suggesting a minimal impact on the Chrysopoidea of the crisis of the diversity at the K-T boundary. Other Cenozoic Chrysopoidea can be attributed to the Chrysopidae sensu stricto.

Long-Proboscid Flies as Pollinators of Cretaceous Gymnosperms

The great evolutionary success of angiosperms has traditionally been explained, in part, by the partnership of these plants with insect pollinators. The main approach to understanding the origins of this pervasive relationship has been study of the pollinators of living cycads, gnetaleans, and basal angiosperms. Among the most morphologically specialized living pollinators are diverse, long-proboscid flies. Early such flies include the brachyceran family Zhangsolvidae, previously known only as compression fossils from the Early Cretaceous of China and Brazil. It belongs to the infraorder Stratiomyomorpha, a group that includes the flower-visiting families Xylomyidae and Stratiomyidae. New zhangsolvid specimens in amber from Spain (ca. 105 mega-annum [Ma]) and Myanmar (100 Ma) reveal a detailed proboscis structure adapted to nectivory. Pollen clumped on a specimen from Spain is Exesipollenites, attributed to a Mesozoic gymnosperm, most likely the Bennettitales. Late Mesozoic scorpionflies with a long proboscis have been proposed as specialized pollinators of various extinct gymnosperms, but pollen has never been observed on or in their bodies. The new discovery is a very rare co-occurrence of pollen with its insect vector and provides substantiating evidence that other long-proboscid Mesozoic insects were gymnosperm pollinators. Evidence is thus now gathering that visitors and probable pollinators of early anthophytes, or seed plants, involved some insects with highly specialized morphological adaptations, which has consequences for interpreting the reproductive modes of Mesozoic gymnosperms and the significance of insect pollination in angiosperm success.

The enigmatic Mesozoic insect taxon Chresmodidae (Polyneoptera): New palaeobiological and phylogenetic data, with the description of a new species from the Lower Cretaceous of Brazil

The morphology of the enigmatic, Mesozoic, aquatic insect family Chresmodidae is redescribed and its phylogenetic affinities among the polyneopterous orders discussed. Study of the complete venation of both fore- and hind wings observed in some specimens from the Spanish Barremian, permit us to postulate the hypothesis that the family belongs to the Archaeorthoptera, thus to the orthopteroid lineage rather than to crown-group Phasmatodea or to the more inclusive group Holophasmatodea (sensu GRIMALDI &E NGEL, 2005). New specimens from Spain, Lebanon, Brazil, and Germany permit a new re-description of some chresmodid body structures with concomitant implications for the phylogenetic position of the family. Chresmoda neotropica n. sp. is described from the Aptian-Albian of the Crato Formation (northeast Brazil). The functional morphology proposed for some of their specialized structures suggest a new hypothesis of Chresmoda palaeo- biology, and related to this some implications for the localized palaeoenvironment as well as global palaeoclimate. The problematic Sternarthron spp. from the Upper Jurassic of Solnhofen were described as probable palpigrades (Arachnida: Palpigradi), based on type material originally thought to be fossil insects. The affinities of Sternarthron HAASE, 1890 have been questioned. Our restudy of HAASEs types clearly confirmed earlier assumptions that these fossils represent nymphal specimens of chresmodids. Consequently, Sternarthron has to be considered as an invalid junior synonym of the fossil insect genera Propygolampis WEYENBERGH, 1874 and Chresmoda GERMAR, 1839.

A rich and diverse tanaidomorphan (Crustacea: Tanaidacea) assemblage associated with Early Cretaceous resin-producing forests in North Iberia: palaeobiological implications

The extinct tanaidomorphan diversity from Early Cretaceous Spanish amber, currently comprising 26 specimens, is reassessed. The fossil family Alavatanaidae Vonk & Schram, 2007, described from Spanish amber, is revised on account of new preparation of type specimens and the discovery of new material. The described tanaidomorphan taxa are classified within the superfamily Paratanaoidea. An emended diagnosis for Alavatanaidae is provided, as well as for the genera Alavatanais Vonk & Schram, 2007 and Proleptochelia Vonk & Schram, 2007, and their respective species Alavatanais carabe Vonk & Schram, 2007 and Proleptochelia tenuissima Vonk & Schram, 2007. Three new species, two of them classified in a new genus each, are described: Alavatanais margulisae Sánchez-García, Peñalver & Delclòs sp. nov., Eurotanais terminator Sánchez-García, Peñalver & Delclòs gen. et sp. nov. and Electrotanais monolithus Sánchez-García, Peñalver & Delclòs gen. et sp. nov. Proleptochelia euskadiensis Vonk & Schram, 2007 is considered a junior synonym of A. carabe, and the genus Proleptochelia, together with its type and only species P. tenuissima, is left without familial placement within Paratanaoidea. Within this superfamily, Alavatanaidae is closely related to Leptocheliidae. Also, morphological variability due to sexual dimorphism in the studied paratanaoids has been determined. Multiple lines of taphonomic and palaeobiological evidence indicate that the Spanish amber tanaids were most likely inhabitants of wet or moist forest floors. http://zoobank.org/urn:lsid:zoobank.org:pub:DC943B18-A01C-412C-8378-C644FEFDA716

Early evolution and ecology of camouflage in insects

Taxa within diverse lineages select and transport exogenous materials for the purposes of camouflage. This adaptive behavior also occurs in insects, most famously in green lacewing larvae who nestle the trash among setigerous cuticular processes, known as trash-carrying, rendering them nearly undetectable to predators and prey, as well as forming a defensive shield. We report an exceptional discovery of a green lacewing larva in Early Cretaceous amber from Spain with specialized cuticular processes forming a dorsal basket that carry a dense trash packet. The trash packet is composed of trichomes of gleicheniacean ferns, which highlight the presence of wildfires in this early forest ecosystem. This discovery provides direct evidence of an early acquisition of a sophisticated behavioral suite in stasis for over 110 million years and an ancient plant–insect interaction.

Primitive Termites in Cretaceous Amber from Spain and Canada (Isoptera)

Abstract The first termites in Early Cretaceous (Albian) amber from Spain are described and figured. Morazatermes krishnai Engel and Delclòs, new genus and species, is described from an imago (and wings of a second specimen) preserved in fossiliferous resin from Moraza, Burgos Province. A second termite species, Cantabritermes simplex Engel and Delclòs, new genus and species, is also recorded from the same deposits but is presently known only from the forewing. Similarly, an isolated forewing in amber from San Just, Teruel Province is described as Aragonitermes teruelensis Engel and Delclòs, new genus and species. Lastly, the first termite in Late Cretaceous (Campanian) amber from Grassy Lake, near Medicine Hat, Canada is described from a fragmentary imago lacking wings or much of the body. All of these taxa belong to a primitive grade of unassigned termites between Mastotermitidae and the families Termopsidae, Hodotermitidae, and Archotermopsidae (sensu Engel et al., 2009). Notes are appended on the recently described “Kalotermes” burmensis Poinar, from the latest Albian of Myanmar (Burmese amber), and the species transferred to Kachinitermopsis Engel and Delclòs, new genus, resulting in the new combination, Kachinitermopsis burmensis. These new taxa highlight the diversity of primitive termites during the Cretaceous.

Long‐proboscid brachyceran flies in Cretaceous amber (Diptera: Stratiomyomorpha: Zhangsolvidae)

The monophyletic family Zhangsolvidae comprises stout‐bodied brachyceran flies with a long proboscis and occurring only in the Cretaceous, originally known in shale from the Early Cretaceous Laiyang Formation (Fm.) in China (Zhangsolva Nagatomi & Yang), subsequently from limestones of the Early Cretaceous Crato Fm. of Brazil. Cratomyoides Wilkommen is synonymized with Cratomyia Mazzarolo & Amorim, both from the Crato Fm.; Cratomyiidae is synonymized with Zhangsolvidae. Two genera and three species of Zhangsolvidae are described: Buccinatormyia magnifica Arillo, Peñalver & Pérez‐de la Fuente, gen. et sp.n. and B. soplaensis Arillo, Peñalver & Pérez‐de la Fuente, sp.n., in Albian amber from Las Peñosas Fm. in Spain; and Linguatormyia teletacta Grimaldi, gen. et sp.n., in Upper Albian–Lower Cenomanian amber from Hukawng Valley in Myanmar. Buccinatormyia soplaensis and Linguatormyia teletacta are unique among all Brachycera, extant or extinct, by their remarkably long, flagellate antennae, about 1.6× the body length in the latter species. A phylogenetic analysis of 52 morphological characters for 35 taxa is presented, 11 taxa being Cretaceous species, which supports placement of the family within Stratiomyomorpha, although not to any particular family within the infraorder.

A New Lineage of Enigmatic Diaprioid Wasps in Cretaceous Amber (Hymenoptera: Diaprioidea)

ABSTRACT A new family of microhymenopteran wasps is described and figured from three new species discovered in Cretaceous amber of Spain (Albian) and New Jersey (Turonian). Spathiopterygidae Engel and Ortega-Blanco, new family, is allied to the Diapriidae and Maamingidae (Proctotrupomorpha: Diaprioidea), sharing with these families putatively derived features relative to Monomachidae. The family contains three genera and three species, all new: Spathiopteryx alavarommopsis Engel and Ortega-Blanco, new genus and species, and Myamaropsis turolensis Engel and Ortega-Blanco, new genus and species, both from the Early Cretaceous (Albian) of Spain, and Spathopria sayrevillensis Engel, Ortega-Blanco, and Grimaldi, new genus and species, from the Late Cretaceous (Turonian) of New Jersey. Spathopria sayrevillensis is reconstructed using x-ray synchrotron microtomography In addition, a peculiar new genus and species, Iberopria perialla Engel, Ortega-Blanco, and Delclòs, of stem-group Diapriidae is described from Spanish amber. The distinctive features and character combinations of these taxa are discussed in connection with possible relationships to the surviving lineages of diaprioids.

A Primitive Aphidiine Wasp in Albian Amber from Spain and a Northern Hemisphere Origin for the Subfamily (Hymenoptera: Braconidae: Aphidiinae)

Abstract A description of a new genus and species of braconid, Archephedrus stolamissus, from Early Cretaceous (Albian) amber from Moraza-Peñacerrada I (Spain) is here provided. This is the first fossil Aphidiinae described in Cretaceous amber. The fossil has some typical characters of the subfamily but possesses a unique assemblage of characters among aphidiines, such as a fairly robust abdomen, with a more pronounced articulation between the first and second, instead of the second and third, metasomal segments, as well as several wing venational traits. The distribution of this and other aphidiine fossils, as well as their putative phylogenetic placement as basal among Aphidiinae, is discussed, supporting a Northern rather than Southern Hemisphere origin for the lineage.