The earliest pollen-loaded insects from the Lower Permian of Russia.

Recent fossil discoveries suggest that the coevolution of insect pollinators and gymnosperms started long before the appearance of flowering plants. One of the keys to understanding the origins of pollination relationships is fossil insects with gymnosperm pollen attached to the body surface. Such fossils are exceedingly rare to find, especially from the Palaeozoic, a time when ambers with insect inclusions were absent. Here, we report compression fossils of Early Permian tillyardembiid insects (Polyneoptera) preserved with pollen on their heads, thoraces, legs and abdomens. This is the earliest finding of pollen-bearing insects, predating the previous oldest record from the Middle Jurassic by ca 120 Ma. Judging by the pollen composition, tillyardembiids visited a narrow range of host plants, including Rufloriaceae (Cordaitales). While it is impossible to say for certain whether tillyardembiids as pollen consumers contributed to pollination, a trophic specialization of this kind could be considered an evolutionary precursor of pollination mutualism.

The first description of an apterous aphid morph of the family Canadaphididae (Hemiptera, Sternorrhyncha, Aphidomorpha).

The first apterous morph of the Canadaphididae family is described based on Canadian amber inclusions. On the basis of morphological features, it was assigned to the species Canadaphis carpenteri. For the first time, a fluorescence microscope was used to examine a representative of the Canadaphididae family, revealing previously unknown details.

Canadaphis mordvilkoi, Kononova 1976 (Hemiptera: Sternorrhyncha: Canadaphididae)redescription and neotype designation.

Here we present a redescription of an aphis species Canadaphis mordvilkoi Kononova, 1976 based on a recently found specimen originating from the same locality as holotype (Yantardakh (Kheta Formation), Taimyr Peninsula, Russia). The new specimen is far better preserved and many characters invisible and not mentioned in original description are available for our analysis. As a result, we were capable to indicate affinities with other species belonging to the Canadaphididae family. As the holotype is destroyed we decided to designate this specimen as a neotype. We also provide the photos of the type genus of this family for the first time.

Possible long-proboscid insect pollinators from the Early Permian of Russia.

Insect pollination is one of the hallmarks of flowering plants.1 Bees, moths, flies, and some other pollinators evolved elongate siphonate mouthparts for sucking concealed nectar and occasionally other liquids.2 However, it is clear from the fossil record that insects with similar adaptations appeared long before the mid-Cretaceous radiation of angiosperms. These insects most probably used their proboscis to reach pollination drops and other sugary fluids that were hidden in the cones of extinct gymnosperms, pollinating them in the process.3-6 The vast majority of these gymnosperm-associated long-proboscid insects have been reported from the Middle Jurassic to the Early Cretaceous, i.e., the time interval that immediately predated the advent of flowering plants.7 By contrast, the Paleozoic stage of the co-evolution between long-proboscid insect pollinators and plants has remained poorly understood. Here, we report a putative pollination mutualism involving long-proboscid holometabolous insects (Panorpida: Protomeropidae) from the Early Permian of Russia (ca. 283-273 Ma). Their elongate mouthparts have very similar morphology to those of some present-day nectarivorous Coleoptera and Hymenoptera and probably served to imbibe micropylar secretions from the semi-closed ovulate organs of the gymnosperms of a peltaspermalean affinity that have been found in the same locality. This is the earliest record of insects with siphonate-like mouthparts, which could indicate that the complex interactions between pollinators and gymnosperms predate the first flowering plants by over 100 Ma.

Morphological and histological study of the forewing of Orthezia urticae (Linnaeus, 1758) (Hemiptera, Sternorrhyncha).

Wings of Orthezia urticae males were studied. Both ventral and dorsal surfaces of wings were examined under light and scanning electron microscopes. The structure regarded as vein cubitus anterior turned out to be a reinforcement element only. Two elements known as radius sector and media are almost transparent depressions in the wing membrane. Veins at the margin of the fold of the wing anal lobe were not confirmed. Studies indicated a row of sensilla cupola at the beginning of the subcostal ridge. Cross sections of the wing membrane showed a two-layered membrane. The presence of two veins was confirmed in a common stem - subcostal and radius. The change of common stem shape was described. Neither tracheae nor nerves were observed. This is the second paper on cross-sections of wing within Sternorrhyncha.

Is there a relationship between the morphology of the forewing axillary sclerites and the way the wing folds in aphids (Aphidomorpha, Sternorrhyncha, Hemiptera)?

The present study describes the relationship between the morphology of the forewing axillary sclerites and the way the wings fold among 24 aphid genera as compared to a representative of coccids. Architecture of the forewing base was imaged with scanning electron and optical (fluorescence) microscopy. Significant differences in morphology of axillary sclerites between aphid species were observed, despite their belonging to one infraorder. Detailed description of 41 features of axillary sclerites was made. There was no difference between axillaries of viviparous (Aphididae) and oviparous (Adelges sp., Phylloxera sp.) species. No clear relationship between morphology of the axillary sclerites and the wing folding could be confirmed. Instead, the thorax structure determines the way the wing folds in aphids. Phylogenetic analysis based on our results cannot be conducted at this stage of study. To show how three-dimensional the structures are and how difficult to describe, a short animation of Aphis fabae (Aphididae) wing base was added. This is a preliminary study about morphology of axillary sclerites among aphids.

New genus and species of the extinct aphid family Szelegiewicziidae and their implications for aphid evolution.

Recently, we are witnessing an increased appreciation for the importance of the fossil record in phylogenetics and testing various evolutionary hypotheses. However, this approach brings many challenges, especially for such a complex group as aphids and requires a thorough morphological analysis of the extinct groups. The extinct aphid family Szelegiewicziidae is supposed to be one of the oviparous lineages in aphid evolution. New material from the rock fossil deposits of Shar Teg (Upper Jurassic of Mongolia), Baissa (Lower Cretaceous of Siberia-Russia), and Burmese amber (Upper Cretaceous of Myanmar) allowed us to undertake a more detailed examination of the morphological features and carry out an analysis of the taxonomical composition and evolution of the family. This led us to the conclusion that evolution of the body plan and wing structure was similar in different, often not closely related groups, probably as a result of convergence. Additionally, we present a description of a new genus and two species (Tinaphis mongolica Zyla &Wegierek, sp. nov., and Feroorbis burmensis Wegierek & Huang, gen. et sp. nov.) that belong to this family.

The forewing of the Aphis fabae (Scopoli 1763) (Hemiptera, Sternorrhyncha): a morphological and histological study.

Dorsal and ventral sides of the forewing of Aphis fabae (Scopoli 1763) (Sternorrhyncha, Hemiptera) were examined by scanning electron microscopy. Reinforcement elements on their surface as well as scale-like elements were described. Using histological methods, cross-sections of the material were done. They showed a two-layered membrane with a circular foramen inside. The course of veins and places of their bifurcation were followed. Common stem of radius (R), media (M), and cubitus anterior (CuA) veins were composed of separate tracheae, which ran separately at the beginning, then continued in a single tunnel, and finally disappeared. Nerves were not observed. Neither were tracheae visible on the further course of those veins. The presence of a devoid-of-trachea costal vein was confirmed. Under scanning electron microscope, convex reinforcements on dorsal side of the wing turned out to be more sclerotized parts of chitin, not giving a zigzag-like profile of the wing on sections. In this paper, we show for the first time a cross-section of a very delicate wing of an aphid representative.

Polyphyly of the extinct family Oviparosiphidae and its implications for inferring aphid evolution (Hemiptera, Sternorrhyncha).

Aphidoidea, the so-called "true aphids" are one of the most challenging groups in terms of solving the phylogenetic relationships. Morphology-based analyses were strongly affected by widespread homoplasy, while the molecular-based attempts struggled with the lack of sufficient phylogenetic signal. Despite significant improvements, the higher classification still remains unresolved and rather controversial. However, the use of the fossil record, one of the most valuable sources of information, was mainly limited to calibration of a phylogenetic tree, without a direct inclusion into the analysis. The extinct family Oviparosiphidae has long been considered as the common ancestor of all recent Aphidoidea and it was used as a calibration point in several analyses, but it has been never analyzed in a phylogenetic context. The family has been treated as a monophyletic group purely based on the simultaneous presence of two abdominal structures, ovipositor and siphunculi. However, it has been shown recently that at least one more extinct lineage, present at the same time, was characterized by the same features. For these reasons, we performed a maximum parsimony analysis using morphological data for extinct aphid taxa to prove the monophyly of Oviparosiphidae. Our analysis shows that the presumed ancestor lineage of recent aphids is a polyphyletic group. Our results support the hypothesis of an early Mesozoic rapid radiation of aphids, which led to several different lineages characterized by both ovipositor and siphunculi. The results indicate the necessity of examining the other extinct families, and shows that the diversity of aphids before the Cretaceous Terrestrial Revolution (KTR) was higher than expected. Even though there is not enough data to perform a formal analysis, fossils seem to suggest a significant impact of the KTR on aphid diversification. Additionally, we have made a redescription of two genera and description of a new species, Vitimaphis subridens sp. nov.

Comparative morphology of the forewing base articulation in Sternorrhyncha compared with a representative of Fulgoromorpha (Insecta, Hemiptera).

The forewing articulation of single species from each of the four subgroups of Sternorrhyncha (Aleyrodomorpha, Aphidomorpha, Coccomorpha, Psyllomorpha) was examined by optical and scanning electron microscopy. The species were compared with a species of Cixiidae (Fulgoromorpha), as an outgroup of Sternorrhyncha. We present the results of a comparative analysis of the forewing articulation in these five groups, propose a standardized terminology and compare our findings with those previously reported. The wing base of all examined species is composed of the following structures: anterior and posterior notal wing process, first, second, and third axillary sclerites, tegula, and axillary cord. The number of elements included in the wing base and the surrounding area is the greatest in Cacopsyllamali, the most complicated species from Sternorrhyncha. Based on the shape of axillary sclerites and the number of elements forming the wing base environment, Orthezia urticae (Coccomorpha) and Cixius nervosus (Fulgoromorpha) are the most similar. Among Sternorrhyncha, the most similar axillaries are those of Aphis fabae and Orthezia urticae, which is congruent with existing classifications. In this paper we show that the four groups from Sternorrhyncha exhibit their own distinct wing base morphology.

The structure of extremely long mouthparts in the aphid genus Stomaphis Walker (Hemiptera: Sternorrhyncha: Aphididae).

Scanning electron microscopy and light microscopy were used to elucidate the morphology of labium and mandibular and maxillary stylets of the aphids Stomaphis quercus (L.) and S. graffii Cholodkovsky. The mechanism of labium shortening associated with feeding process was described as well. Stomaphis quercus and S. graffii have cone-shaped labium of 13 and 10 mm in length, respectively, that strongly extend behind the abdomen. The stylets bundle comprises a pair of mandibular and maxillary stylets which are on average as long as labium. Serial cross sections of labium revealed that the first segment is inverted inside and the second is pulled into it; both segments are shifted into abdomen. This study provides new information on S. quercus and S. graffii mouthparts that may help to understand their feeding behavior.

New mayfly genera from the Middle Triassic of Poland and their evolutionary and paleogeographic implications (Ephemerida: Litophlebiidae, Vogesonymphidae).

Two new mayfly genera and species from the Triassic deposits of the Palegi area (southeast Poland) are described. This is the first description of aquatic insects from the Palegi locality. Triassolitophlebia palegica gen. et sp. nov. (Litophlebiidae) is established on the basis of an isolated forewing. This is the first finding of this family in the Northern Hemisphere, known previously only from the Molteno Formation (South Africa). This is also the first mayfly family from the Triassic which has been found in both Hemispheres, providing additional evidence of the presumed similarity of aquatic insect faunas in the Southern and Northern Hemispheres during the Triassic. The consistent wing venation of ancient mayflies with homonomous wings could be evidence that they originated from the same ancestor. The second new mayfly, Palegonympha triassica gen. et sp. nov. (Vogesonymphidae), is described on the basis of a single fossil nymph (imprint of the exuviae) and indicates the similarity of the Palegi arthropod assemblage to that described from the Middle Triassic of France. The presence of a mayfly nymph in the last instar stage suggests not only that the Palegi deposit represents a fluvial environment with well-oxygenated and limpid water but also that these conditions lasted long enough to allow for such development.

Revised concept of the fossil genus Oviparosiphum Shaposhnikov, 1979 with the description of a new genus (Hemiptera, Sternorrhyncha, Aphidomorpha).

This paper presents a revision of the aphid genus Oviparosiphum, which is known from the Cretaceous period. Redescriptions of two species: Oviparosiphumjakovlevi Shaposhnikov, 1979 and Oviparosiphumbaissense Shaposhnikov & Wegierek, 1989 are made, and an updated diagnosis of this genus is provided. Oviparosiphumbaissense is the type species of a newly described genus Archeoviparosiphum gen. n. Five other species of Oviparosiphum are also transferred to the new genus. The basis for their separation from Oviparosiphum is the structure of the siphunculi and ovipositor. A key is provided to the genera of Oviparosiphidae.

Chauliodites niedzwiedzkii sp. n. (Grylloblattida: Chaulioditidae) from Triassic sediments of Poland.

A new representative of the family Chaulioditidae (Insecta, Grylloblattida), Chauliodites niedzwiedzkii sp. n., is described from the Upper Olenekian-Lower Anisian sediments of Palegi in Holy Cross Mountains, Poland. This is the first formal description of any fossil insect from Palegi area.

A new family of aphids (Hemiptera, Sternorrhyncha) from the Lower Cretaceous of Baissa, Transbaikalia.

The family Rasnitsynaphididaefam. n. has a unique combination of characters: 9-segmented antennae; rhinaria arranged in many transverse rows, surrounding the antennal segments; segment IX narrower than other segments of flagellum, always without rhinaria; cubitus branches separated; ovipositor present; siphuncular pores absent. The new family comprises the genus Rasnitsynaphisgen. n. with three species, Rasnitsynaphis ennearticulatasp. n., Rasnitsynaphis coniunctasp. n., and Rasnitsynaphis quadratasp. n., all from the Lower Cretaceous of Transbaikalia.