Antonio Arillo

A NEW FOSSIL RESIN WITH BIOLOGICAL INCLUSIONS IN LOWER CRETACEOUS DEPOSITS FROM ÁLAVA (NORTHERN SPAIN, BASQUE-CANTABRIAN BASIN)

Abstract The occurrence of amber in Sierra de Cantabria (Álava, Basque Country) has been known for more than two decades but biological inclusions have only recently been found. The existence of crustaceans (amphipods and isopods), chelicerates (acari and arachnids), 12 orders of insects, and several bird feathers are reported in this preliminary study. In addition, there are leaf remains, molluscs, and a fair number of inorganic inclusions. Pollen analysis of the clastic series indicates an age between upper Aptian—middle Albian, which allows an assignment of this stratigraphic unit to the Nograro Formation. Chemical analysis indicates that the amber has high maturity, which reflects its Cretaceous age. Chemical composition analysis also indicates an araucariacean origin, which is corroborated by pollen found within the amber deposit. This new fossil site provides information for the reconstruction of paleocommunities of arthropods and sedimentary environments in the extreme south of the Basque-Cantabrian Basin during the Lower Cretaceous, characterized by coniferous forests with an understory of vascular cryptograms. Some of the identified arthropods add to the fossil record for various groups that are poorly known or unknown for this time period. This Lagerstätte constitutes one of the most important deposits of Mesozoic amber in the world.

Brachyceran Diptera (Insecta) in Cretaceous ambers, Part IV, Significant New Orthorrhaphous Taxa

Abstract Thirteen species of basal Brachycera (11 described as new) are reported, belonging to nine families and three infraorders. They are preserved in amber from the Early Cretaceous (Neocomian) of Lebanon, Albian of northern Spain, upper Albian to lower Cenomanian of northern Myanmar, and Late Cretaceous of New Jersey USA (Turonian) and Alberta, Canada (Campanian). Taxa are as follows, with significance as noted: In Stratiomyomorpha: Stratiomyidae (Cretaceogaster pygmaeus Teskey [2 new specimens in Canadian amber], Lysistrata emerita Grimaldi & Arillo, gen. et sp. n. [stem-group species of the family in Spanish amber]), and Xylomyidae (Cretoxyla azari Grimaldi & Cumming, gen. et sp. n. [in Lebanese amber], and an undescribed species from Spain). In Tabanomorpha: Tabanidae (Cratotabanus newjerseyensis Grimaldi, sp. n., in New Jersey amber). In Muscomorpha: Acroceridae (Schlingeromyia minuta Grimaldi & Hauser, gen. et sp. n. and Burmacyrtus rusmithi Grimaldi & Hauser gen. et sp. n., in Burmese amber, the only definitive species of the family from the Cretaceous); Mythicomyiidae (Microburmyia analvena Grimaldi & Cumming gen. et sp. n. and Microburmyia veanalvena Grimaldi & Cumming, sp. n., stem-group species of the family, both in Burmese amber); Apsilocephalidae or near (therevoid family-group) (Kumaromyia burmitica Grimaldi & Hauser, gen. et sp. n. [in Burmese amber]); Apystomyiidae (Hilarimorphites burmanica Grimaldi & Cumming, sp. n. [in Burmese amber], whose closest relatives are from the Late Jurassic of Kazachstan, the Late Cretaceous of New Jersey, and Recent of California). Lastly, two species belonging to families incertae sedis, both in Burmese amber: Tethepomyiidae (Tethepomyia zigrasi Grimaldi & Arillo sp. n., the aculeate oviscapt of which indicates this family was probably parasitoidal and related to Eremochaetidae); and unplaced to family is Myanmyia asteiformia Grimaldi, gen. et sp. n., a minute fly with highly reduced venation. These new taxa significantly expand the Mesozoic fossil record of rare and phylogenetically significant taxa of lower Brachycera.

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.

Paleoethology: fossilized behaviours in amber

When organisms became trapped in sticky resin they usually expired so quickly that they could fossilize in almost lifelike circumstances. Based on frequent description in literature of many examples we propose in this paper a classification of these paleobehaviours.

Did dinosaurs have any relation with dung‐beetles? (The origin of coprophagy)

It is widely accepted that Mesozoic ecosystems were basically similar to Cenozoic ecosystems and it has been proposed that the role of dung‐beetles in those ecosystems was identical to that of today, but the dung of dinosaurs were used as a source of food instead of the dung of mammals. While dinosaurs have been known since Triassic, Scarabeids are present in the fossil records probably since Lower Jurassic. But a very important metabolic feature of dinosaurs has not been taken into account, the connection between digestive and uro‐genital systems. So here we propose the hypothesis that coprophagy in dung‐beetles has been associated, since it began, with mammals.

Description of a new hypogean species of the genus Trechus Clairville, 1806 from eastern Spain and comments on the Trechus martinezi‐lineage (Coleoptera: Adephaga: Carabidae)

Trechus torressalai new species, a cave‐dwelling ground beetle from eastern Spain is described. Ecological data are also included. Diverse morphological characters suggest they belong in the Trechus fulvus‐group. Detailed study of the aedeagus suggests taxonomic affinities with Trechus martinezi Jeannel, Trechus alicantinus Español, and Trechus beltrani Toribio. The close relationship among these species reveals an adelphotaxon‐complex that constitutes the T. martinezi‐lineage, exclusive to the north‐eastern Betic Mountains. An identification key to the species and biogeographical comments on the T. martinezi‐lineage are also included. Se describe Trechus torressalai nueva especie, carábido cavernícola del Este de España. Se incluyen datos sobre su ecología. Diversos caracteres morfológicos aconsejan su inclusión dentro del amplio grupo de Trechus fulvus. El estudio detallado de la genitalia masculina sugiere su proximidad taxonómica a Trechus martinezi Jeannel, Trechus alicantinus Español y Trechus beltrani Toribio. Las estrechas afinidades entre estas cuatro especies desvela un complejo de adelfotaxones que constituyen el linaje de T. martinezi, exclusivo del extremo nororiental de las montañas Béticas. Se aportan claves de identificación para estas especies. Se incluyen comentarios sobre la biogeografía del linaje de T. martinezi.

A Review of the Eurasian fossil species of the bee Apis

Fossil Apis species from the Oligocene, Miocene and Plio-Pleistocene of Eurasia are described and their relationships with Recent species are discussed. Several new populations of fossil bees are reported from the Oligocene and Miocene of France and Spain, including Apis aquisextusensis sp. nov. The present state of knowledge of fossil bee systematics is poor because of the general lack of preserved characters. Some of the problems, and items requiring further investigation, are identified.

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 species of fossil oribatid mite (Acariformes, Oribatida, Trhypochthoniidae) from the Lower Cretaceous amber of San Just (Teruel Province, Spain)

Abstract A new fossil species, Trhypochthonius lopezvallei sp. nov. (Trhypochthoniidae), is described based on one specimen preserved in amber from the San Just outcrop (Teruel Province, Spain) believed to be Albian in age. A comparison with Recent and fossil Trhypochthoniidae is given. A new name, Sachalinbates, is proposed to replace Sachalinella (a fossil oribatid genus described from Sakhalin Paleocene amber) which is preoccupied.

The first Baltic amber dysagrionine damselfly (Odonata: Zygoptera: Thaumatoneuridae: Dysagrioninae)

Abstract A new genus and species Electrophenacolestes serafini is described. It is the first Thaumatoneuridae recorded from an amber deposit and the second record of the family in the European Paleogene. A comparison with related genera and families is done.