The male postabdomen and genital apparatus of †Mengea tertiaria,a strepsipteran amber fossil (Insecta)

The male genital apparatus of a fossil insect including internal soft parts is described in detail for the first time. The conditions found in an approximately 42‐My‐old specimen of ?Mengea tertiaria embedded in Baltic amber are compared to what is found in other extinct and extant strepsipterans, notably members of the basal ‘Mengenillidae’ (probably paraphyletic). The postabdomen of ?Mengea is very similar to what is present in other fossil and extant members of Strepsiptera. Only few structural features vary within the group, but it differs strongly from the apparatus in other holometabolan lineages. The slender, exposed tergite IX, the complete absence of parameres, a sperm pump formed by a strongly developed muscularis (layers of mainly circular muscle fibres) around the proximal part of the ejaculatory duct and the presence of four specific muscles are potential autapomorphies of the Strepsiptera. A presumptive strepsipteran ground plan feature found in ?Mengea is the nearly straight, simple penis, which is also present in ?Protoxenos, ?Cretostylops, Bahiaxenos, Mengenilla, Eoxenos and Congoxenos. This strongly suggests that males of ?Mengea (and other stem group strepsipterans) copulated in a very similar way as males of extant members of the group with free‐living females (e.g. Mengenilla). In contrast, the penis of stylopidian males, which copulate with females parasitizing in pterygote hosts, is hook shaped. A sister group relationship between ?Mengea and Strepsiptera s.s. (extant groups) is supported by a distinctly weaker sclerotization of the abdominal tergites, compared to the corresponding sternites. The study of other stem group strepsipterans using μ‐computer tomography should have high priority. This technique has a great potential to facilitate morphological reconstruction and phylogenetic placement of amber fossils.     

Phylogeny of the Strepsiptera based on morphological data of the first instar larvae

This investigation was the first cladistic analysis using morphological data of first instar larvae of Strepsiptera. The analysis of representatives of nearly all known families of Strepsiptera supports the division of Strepsiptera into Mengenillidia and Stylopidia. Corioxenidae and Elenchidae are placed at the base of Stylopidia. Halictophagidae is the sister group to Xeninae + Myrmecolacidae + Stylopinae. Xeninae is placed as the sister group to Myrmecolacidae + Stylopinae. Stylopidae are paraphyletic. Thus, Xenidae stat. n. is re-established. A sister-group relationship between Myrmecolacidae and Elenchidae is not supported on characters of first instar larvae.     

†Kinzelbachilla ellenbergeri – a new ancestral species,genus and family of Strepsiptera (Insecta)

A single male specimen of a new species (†Kinzelbachilla ellenbergeri gen. et sp.n. ) of a new family of the endoparasitic Strepsiptera (†Kinzelbachillidae fam.n.) from Burmese amber is described and evaluated with respect to its systematic placement. Its features come very close to the presumptive groundplan of the order suggested in recent studies. Preserved plesiomorphic features are the following: fully sclerotized head with long coronal suture, small ommatidia not separated by chitinous bridges, absence of microtrichia between ommatidia, eight antennomeres, robust mandibles with dicondylic articulation, galea distinctly developed, free pro‐ and mesotrochanters, slender five‐segmented tarsi without adhesive soles, and equally sclerotized abdominal tergites and sternites. An important character that is not recognizable due to damage is the shape of the metapostnotum. This structure is transverse in the groundplan of Strepsiptera and in †Protoxenos, but elongated and shield‐like in all other known strepsipterans. In a cladistic analyses of 82 characters of adult males and additional characters for females and immatures (scored as unknown for all included fossils) †Kinzelbachilla is placed as sister group of all remaining strepsipterans except for †Protoxenos, followed by †Cretostylops and †Mengea as the third and fourth branches in the stem group, respectively. The position of †Protoxenos as first branch is suggested by three unambiguous apomorphic features of all remaining Strepsiptera, the reduced size of less than 6 mm, mandibles distinctly narrowing distad the basalmost part, and fan‐shaped hindwings which are broader than they are long. The hitherto known fossil stem group strepsipterans do not distinctly narrow the large morphological gap separating this order from its sister group, the Coleoptera. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:07554C01-DEC3-4080-A337-B1F46BC9070F .     

A case of mistaken identification of an amber fossil male strepsipteran (Insecta: Strepsiptera)

A well-preserved specimen of amber Strepsiptera was erroneously described as Stylops neotropicallis Kogan and Poinar, 2010. The taxonomic position of the species was based on a count of six antennomeres (typical of the Stylopidae); however, further observations showed that there are actually seven antennomeres, which places the fossil in the family Myrmecolacidae, and it is herein redescribed in the genus Palaeomyrmecolax Kulicka, 2001. Furthermore, nuclear magnetic resonance spectroscopy analysis of a sample of the amber piece containing the fossil revealed that it was Baltic rather than Dominican amber as originally thought. The fossil was compared with another specimen from Baltic amber in the Poinar collection, which shows close affinity to Palaeomyrmecolax succineus Kulicka, 2001, the type species of the genus. Palaeomyrmecolax neotropicallis (n.comb.) differs from that specimen and seems to differ also from the other four species in the genus Palaeomyrmecolax.     

The evolution of Strepsiptera (Hexapoda)

An evolutionary scenario for the enigmatic group Strepsiptera is provided, based on the results of a comprehensive cladistic analysis of characters of all life stages. A recently described fossil--+Protoxenos janzeni--the most archaic strepsipteran, sheds new light on the early evolution of the group and reduces the "morphological gap" between Strepsiptera and other insects. It weakens both current hypotheses--Coleoptera+Strepsiptera and Diptera+Strepsiptera (="Halteria"). The splitting into +Protoxenos (Protoxenidae) and the remaining Strepsiptera was linked with a distinct size reduction and many morphological changes. Unlike males of extant strepsipteran species, +Protoxenos was still able to process food. Mengeidae (+Mengea), with two small species, is the sister group of extant Strepsiptera. A unique characteristic of extant males (Strepsiptera s. str.) is the mouthfield sclerite. It is part of an air uptake apparatus which belongs to an extremely modified air-filled "balloon gut". Besides this, male strepsipterans possess specialised antennae and compound eyes, a strongly developed flight apparatus, large testes, and a sperm pump, whereas other organ systems are strongly reduced (e.g., fat body, malpighian tubules). Males are designed to find females within a few hours and to copulate. A dramatic change is linked with the split into Mengenillidae and Stylopidia. The change to pterygote hosts and the permanent endoparasitism of the females are evolutionary novelties acquired by the latter clade, and linked with far-reaching morphological transformations, e.g. the presence of unique brood organs. Hairy tarsal adhesive devices are present in males and guarantee efficient attachment to the host during copulation. A well-founded clade within Stylopidia is Stylopiformia, which are characterised by a unique fissure-shaped birth opening. The evolutionary development towards the most specialised and successful forms (parasites of aculeate Hymenoptera [e.g., Xenidae+Stylopidae], ca. 46% of the species) is a stepwise process. The presented evolutionary scenario comprises a complex network of functionally correlated morphological changes in primary larvae, secondary larvae, females and males.     

Protoxenidae fam. nov. (Insecta, Strepsiptera) from Baltic amber — a 'missing link' in strepsipteran phylogeny

A male specimen of a new strepsipteran genus and species ( Protoxenos janzeni gen. et sp. nov.) and family (Protoxenidae fam. nov.) found in Baltic amber is described and illustrated. It shows features which are apparently more plesiomorphic than in hitherto known strepsipterans, such as laterally inserted eight-segmented antennae, very robust mandibles with a broad base, a prominent galea, a comparatively short, transverse metapostnotum, hindwings that are feebly extended in a rostrocaudal direction, and equally sclerotized abdominal tergites and sternites. Based on a cladistic analysis of 46 characters of males of 11 genera and three outgroup taxa, P. janzeni is the sister group of all other known strepsipterans, and Mengea the sister group of Strepsiptera s.s . Eoxenos is the sister group of the remaining extant strepsipterans and Mengenillidae is therefore paraphyletic. Newly established groundplan features of Strepsiptera will facilitate the clarification of the systematic position of the Order in future studies.     

Sciadopitys cladodes from Eocene Baltic amber

The Baltic amber deposit represents the largest accumulation of any fossil resin worldwide and hundreds of thousands of entrapped arthropods have been recovered so far. The source plants of Baltic amber, however, are still controversial, and the botanical composition of the ‘Baltic amber forest’ remains poorly studied. Here, we provide the first unequivocal Baltic amber inclusions of the umbrella pine Sciadopitys (Sciadopityaceae), a genus that has been suggested as the source of succinite (the main variety of Baltic amber) based on chemical analyses. As previously suggested sciadopitoid inclusions must be reconsidered as being notional, representing angiosperm leaves instead, the new fossils are the first unambiguous macrofossil evidence of Sciadopitys from the ‘Baltic amber forest’, and the first pre‐Oligocene macrofossil record of Sciadopitys from Europe. The fossil Sciadopitys cladodes provide new insights into the conifer diversity of the ‘Baltic amber forest’ and broaden the picture of its palaeoecology, indicating the presence of humid swamp to raised bog habitats. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180 , 258–268.     

The abdomen of a free-living female of Strepsiptera and the evolution of the birth organs

Mengenillidae is a small, basal family of Strepsiptera, mainly characterized by free-living females in contrast to the endoparasitic females of Stylopidia. Here, we describe external and internal structures of the female abdomen of Eoxenos laboulbenei (Mengenillidae). The external morphology was examined and documented using microphotography. Internal structures were reconstructed three-dimensionally using a μCT-data set. The morphologically simplified abdomen comprises 10 segments. The integument is weakly sclerotized and flexible. Spiracles are present dorsolaterally on segments I–VII. Segment VII bears the posteroventral birth opening and the small abdominal segment X carries the anus at its apex. Numerous eggs float freely in the hemolymph. The musculature of segments I–IV is composed of ventral and dorsal longitudinal muscle bundles, strongly developed paramedial dorsoventral muscles and a complex meshwork of small pleural muscles, with minimal differences between the segments. Segments V–X contain more than 50 individual muscles, even though the musculature as a whole is weakly developed. Even though it is not involved in processing food, the digestive tract is well-developed. Its postabdominal section comprises a part of the midgut and the short hindgut. The midgut fills a large part of the postabdominal lumen. The lumina of the midgut and hindgut are not connected. Five or six nodular Malpighian tubules open into the digestive tract at the border region between the midgut and hindgut. The birth organ below the midgut releases the primary larvae after hatching via the birth opening at segment VII. It is likely derived from primary female genital ducts. The presence of six additional birth organs of segments I–VI are de novo formations and a groundplan apomorphy of Stylopidia, the large strepsipteran subgroup with endoparasitic females. The loss of the primary birth organ of segment VII is an apomorphy of Stylopiformia (Stylopidia excl. Corioxenidae).     

How diverse were ferns in the Baltic amber forest?

Diverse temperate forest types and a high atmospheric humidity have recently been suggested for the Eocene source area of Baltic amber. However, ferns are astonishingly rare as inclusions in this amber, which is in contrast to other seed‐free land plants, fungi, and lichens. Moreover, the identities of some of the few described putative fern taxa are dubious, and some fossils were even assigned to the Paleozoic seed fern genera Alethopteris, Pecopteris and to the form genus Sphenopteris containing Paleozoic and Mesozoic fern‐like leaf fossils. Here, we review previously described fern inclusions from Baltic amber and identify further fern‐like leaf inclusions as belonging to the extant angiosperm genus Comptonia (sweet ferns, Myricaceae). We conclude that only one taxon, Matonia striata (Matoniaceae), can with confidence be identified as a Polypodiopsida representative. Although “Pecopteris” humboldtiana is so far only known as sterile foliage, its leaf morphology strongly suggests that also this taxon belongs to the Polypodiopsida rather than to any other tracheophyte lineage. We propose accommodating “Pecopteris” humboldtiana in the new genus Berendtiopteris. “Alethopteris” serrata and “Sphenopteris” phyllocladoides are not to be regarded as evidence of ferns from Baltic amber. Reinvestigation of the holotypes of these two taxa did not reveal to which tracheophyte lineages these fossils belong. We suggest that the scarcity of fern remains from Baltic amber may reflect both a relatively low fern diversity in the source area of the fossil resin, and an absence or rarity of epiphytic and climbing ferns as observed in modern temperate forest ecosystems.     

Description of a new genus and two new species of Corioxenidae from Jordan (Insecta: Strepsiptera)

Abstract

Two new species of twisted-wing insects (Strepsiptera) of the family Corioxenidae, Proceroxenos jordanicus n.gen., n.sp., and Dundoxenos breviphlebos n.sp., are described from Jordan. Identification keys to the subfamilies of the Corioxenidae, the genera of the subfamily Uniclavinae, and the species of the genus Dundoxenos are provided. A list of all described species of Corioxenidae is given, including information on geographic distribution, and host species when available.     

Phylogenetic position of the Strepsiptera: Review of molecular and morphological evidence

Molecular evidence of the monophyly of the Halteria (Strepsiptera + Diptera) is reviewed. The majority of morphological characters, which have classically been used to establish a Strepsiptera + Coleoptera sister group, are rejected, because they are based on erroneous interpretations of strepsipteran morphology. The scorings of 31 morphological characters, which directly relate to the phylogenetic position of Strepsiptera, are provided, and their distribution and optimization on the molecular + morphological tree is discussed. Of these, 13 characters specifically support the placement of Strepsiptera within the Mecopterida; seven of which are based on the optimization of inapplicable or missing data, and six of which are based on states that can be scored for Strepsiptera. Only a single character (posteromotorism) suggests a sister group relationship with the Coleoptera. The morphological and molecular data are largely congruent, and suggest that the Strepsiptera are sister group to the Diptera.     

The Strepsiptera problem: phylogeny of the holometabolous insect orders inferred from 18S and 28S ribosomal DNA sequences and morphology

Phylogenetic relationships among the holometabolous insect orders were inferred from cladistic analysis of nucleotide sequences of 18S ribosomal DNA (rDNA) (85 exemplars) and 28S rDNA (52 exemplars) and morphological characters. Exemplar outgroup taxa were Collembola (1 sequence), Archaeognatha (1), Ephemerida (1), Odonata (2), Plecoptera (2), Blattodea (1), Mantodea (1), Dermaptera (1), Orthoptera (1), Phasmatodea (1), Embioptera (1), Psocoptera (1), Phthiraptera (1), Hemiptera (4), and Thysanoptera (1). Exemplar ingroup taxa were Coleoptera: Archostemata (1), Adephaga (2), and Polyphaga (7); Megaloptera (1); Raphidioptera (1); Neuroptera (sensu stricto = Planipennia): Mantispoidea (2), Hemerobioidea (2), and Myrmeleontoidea (2); Hymenoptera: Symphyta (4) and Apocrita (19); Trichoptera: Hydropsychoidea (1) and Limnephiloidea (2); Lepidoptera: Ditrysia (3); Siphonaptera: Pulicoidea (1) and Ceratophylloidea (2); Mecoptera: Meropeidae (1), Boreidae (1), Panorpidae (1), and Bittacidae (2); Diptera: Nematocera (1), Brachycera (2), and Cyclorrhapha (1); and Strepsiptera: Corioxenidae (1), Myrmecolacidae (1), Elenchidae (1), and Stylopidae (3). We analyzed approximately 1 kilobase of 18S rDNA, starting 398 nucleotides downstream of the 5' end, and approximately 400 bp of 28S rDNA in expansion segment D3. Multiple alignment of the 18S and 28S sequences resulted in 1,116 nucleotide positions with 24 insert regions and 398 positions with 14 insert regions, respectively. All Strepsiptera and Neuroptera have large insert regions in 18S and 28S. The secondary structure of 18S insert 23 is composed of long stems that are GC rich in the basal Strepsiptera and AT rich in the more derived Strepsiptera. A matrix of 176 morphological characters was analyzed for holometabolous orders. Incongruence length difference tests indicate that the 28S + morphological data sets are incongruent but that 28S + 18S, 18S + morphology, and 28S + 18S + morphology fail to reject the hypothesis of congruence. Phylogenetic trees were generated by parsimony analysis, and clade robustness was evaluated by branch length, Bremer support, percentage of extra steps required to force paraphyly, and sensitivity analysis using the following parameters: gap weights, morphological character weights, methods of data set combination, removal of key taxa, and alignment region. The following are monophyletic under most or all combinations of parameter values: Holometabola, Polyphaga, Megaloptera + Raphidioptera, Neuroptera, Hymenoptera, Trichoptera, Lepidoptera, Amphiesmenoptera (Trichoptera + Lepidoptera), Siphonaptera, Siphonaptera + Mecoptera, Strepsiptera, Diptera, and Strepsiptera + Diptera (Halteria). Antliophora (Mecoptera + Diptera + Siphonaptera + Strepsiptera), Mecopterida (Antliophora + Amphiesmenoptera), and Hymenoptera + Mecopterida are supported in the majority of total evidence analyses. Mecoptera may be paraphyletic because Boreus is often placed as sister group to the fleas; hence, Siphonaptera may be subordinate within Mecoptera. The 18S sequences for Priacma (Coleoptera: Archostemata), Colpocaccus (Coleoptera: Adephaga), Agulla (Raphidioptera), and Corydalus (Megaloptera) are nearly identical, and Neuropterida are monophyletic only when those two beetle sequences are removed from the analysis. Coleoptera are therefore paraphyletic under almost all combinations of parameter values. Halteria and Amphiesmenoptera have high Bremer support values and long branch lengths. The data do not support placement of Strepsiptera outside of Holometabola nor as sister group to Coleoptera. We reject the notion that the monophyly of Halteria is due to long branch attraction because Strepsiptera and Diptera do not have the longest branches and there is phylogenetic congruence between molecules, across the entire parameter space, and between morphological and molecular data.     

Bahiaxenidae, a "living fossil" and a new family of Strepsiptera (Hexapoda) discovered in Brazil

An adult male of a newly discovered strepsipteran species from Brazil— Bahiaxenos relictus— is described. A new family Bahiaxenidae is suggested based on cladistic analyses of comprehensive morphological data sets with a broad taxon sampling including the stem group. It is unambiguously placed as the sister group of all other extant families of Strepsiptera. Bahiaxenos relictus is the only species of basal, i.e. non-stylopidian, Strepsiptera occurring in the New World. It appears to be a relict taxon that has survived in the fossil sand dunes of the São Francisco River (Bahia State). The loss of the 8th antennomere and the greatly reduced labrum are autapomorphies of Strepsiptera s.s . excluding Bahiaxenidae. The sister group relationship between†Protoxenidae and the remaining Strepsiptera, and between † Cretostylops and a clade comprising † Mengea and Strepsiptera s.s. , is confirmed, as is the monophyly of Stylopidia and Stylopiformia.     

Olemehlia krali,a new genus and species of Ripidiinae from Baltic amber (Coleoptera: Ripiphoridae)

A new genus and species, Olemehlia krali n. g. & n. sp. (Coleoptera: Ripiphoridae: Ripidiinae), is described from Baltic amber. The newly described genus is compared with all known fossil and extant closely related genera of the subfamily Ripidiinae. It is characterised by the following combination of characters: antennae consisting of 10 antennomeres, with scape, pedicel, and antennomere 3 simple and antennomeres 4–10 uniflabellate, mouthparts reduced to a small single unpaired tubercle, postocular ommatidia absent, pronotum trapezoid without grooves or impressions, elytra without white apical spots, tarsal formula 5–5–4, hind tarsomere I shorter than three remaining combined. Diversity of Ripiphoridae in Baltic amber is discussed; key to Ripidiinae genera from Baltic amber is provided.     

Using fossils to determine an amber source: aphids and crane flies in Chinese or Baltic amber?

Aphids and crane flies in amber purported to have originated in China were examined. Six aphid specimens were characterised and determined to be conspecific with Germaraphis defuncta Heie, originally described from Baltic amber. Two crane flies from the same collection were identified as Cheilotrichia minuta and Rhabdomastix brevis, species also previously described from Baltic amber. This study shows that when the locality of an amber source is ambiguous, fossils can be used to determine the true province.     

Diplocoelus probiphyllus n. sp., the first known fossil False Skin Beetle (Coleoptera: Biphyllidae)

The first fossil species of the family Biphyllidae LeConte 1861, Diplocoelus probiphyllus n. sp., is described from Baltic amber. This species, which shows intermediate characters of antennae, sculpturing and pubescence between Diplocoelus Guérin-Ménéville 1844 and Biphyllus Dejean 1821, also suggests the presence of temperate mixed forests and a climate cooler than it has been until today hypothesised for Baltic Amber.     

The sperm pumps of Strepsiptera and Antliophora (Hexapoda)

Male genital structures of representatives of Strepsiptera, Siphonaptera and Diptera are described in detail, with special emphasis on sperm pumps. The parts involved in the apparatus are evaluated with respect to their homology. Functional interpretations are presented based on the morphological observations. The phylogenetic significance of characters related to the male genital apparatus is discussed. The sperm pumps differ strongly in Strepsiptera and Antliophora (s.s.) and are not homologous. The strepsipteran type, which lacks any sclerotized parts, has evolved independently. Autapomophies of the male genital apparatus are the compact testes, the large balloon‐shaped vesicula seminalis, the strongly developed musculature of the proximal ductus ejaculatorius, the strongly simplified copulatory organ, the unusual muscles of segments VIII and IX, and the complete absence of accessory glands. The median fusion and almost globular shape of the vesicula are potential autapomorphies of Corioxenidae. The absence of the furrow separating the testes from the vesicula seminalis is a derived condition found in Xenos and Myrmecolax. A sperm pump is absent in Boreus (Mecoptera) and Culicomorpha and the functionally relevant parts and their arrangement differ strongly in Siphonaptera, Pistillifera and Diptera (excl. Culicomorpha). The presence of a functional and homologous pumping apparatus does not belong to the groundplan of Antliophora, which implies that this alleged autapomorphy of the clade is invalid. A sperm pump belongs to the groundplan of Diptera and was secondarily reduced in Culicomorpha, many representatives of Bibionomorpha, and in Diopsidae. It was very likely primarily absent in Mecoptera. However, the precise reconstruction of the groundplan depends on the position of Nannochoristidae within Mecoptera and on the possible affinities of Siphonaptera and Boreidae. Sperm pumps should be considered as a functional term and not be used as a character for phylogenetic reconstruction, unless specific similarities are included in the character definition.     

Phylogeny of ensign scale insects (Hemiptera: Coccoidea: Ortheziidae) based on the morphology of Recent and fossil females

The Ortheziidae (ensign scale insects) is a morphologically well‐defined family. The morphology and occurrence in the fossil record suggests a probable early origin of the family in scale insect evolution. The present phylogenetic analysis – based on 69 morphological characters of female ortheziids, using 39 exemplar Recent species – provides the first analytical assessment of relationships among living genera of the family, as well as the relationships of eight fossil species, based on complete, well‐preserved specimens in amber. Monophyly of the subfamilies Newsteadiinae, Ortheziinae and Ortheziolinae is supported, but Nipponortheziinae is found to be paraphyletic by inclusion of the Ortheziolinae. Thus, the subfamily Ortheziolinae is reduced in rank to tribe Ortheziolini stat.n. , which now includes Matileortheziola Kozár & Foldi, Ortheziolacoccus Kozár, Ortheziolamameti Kozár and Ortheziola?ulc. Consequently, the tribes Matileortheziolini, Ortheziolacoccini and Ortheziolamametini are synonymized ( syn.n. ) here under Ortheziolini. Five new species and one new genus of fossil ensign scales are described from three amber deposits: Burmorthezia gen.n. with type species Burmorthezia kotejai sp.n. and also B. insolita sp.n ., both in mid‐Cretaceous Burmese amber (98 Ma) and Arctorthezia baltica sp.n. in Eocene Baltic amber (c. 43 Ma) based on second‐instar nymphs; Mixorthezia kozari sp.n . and M. dominicana sp.n . in Miocene Dominican amber (c. 17 Ma) based on adult females. Fossil placements are unambiguous, with Burmorthezia forming a stem to crown‐group (Recent and Tertiary) Ortheziidae. A summary of described fossil ortheziids is provided.     

Hepatics from Rovno amber (Ukraine), 6. Frullania rovnoi,sp. nov.

A fossil species of the extant liverwort genus Frullania Raddi is described and illustrated, based on a single inclusion in a piece of Rovno amber (Ukraine) that shares its age with Late Eocene Baltic amber, its northern contemporary. Frullania rovnoi is characterised by leaves with a rounded dorsal lobe and the absence of ocelli. The ventral lobe is inflated and forms a saclike lobule, which is bell-shaped and somewhat constricted above the mouth. The bifid underleaves have several blunt teeth or angulations along the shoulder. The Rovno fossil differs sufficiently from morphologically similar species preserved in Baltic and Bitterfeld amber as to be described as new to science. The shape of the lobules and underleaves, as well as the absence of ocelli, indicate an affiliation to F. sect. Australes, hitherto represented in Eocene amber inclusions solely by F. schumannii (Casp.) Grolle. The Rovno fossil is distinguished from extant species of F. subg. Australes and from F. schumannii by having roughly and irregularly dentate-angulate underleaf margins.     

Have ladybird beetles and whiteflies co-existed for at least 40 Mya?

The beetle family Coccinellidae is rarely recorded from fossils. Most of the records come from the nineteenth and beginning of the twentieth century. A complete list of Coccinellidae records from Baltic amber is presented and discussed. Extensive literature research provided a surprising conclusion that not a single species of Coccinellidae has been formally described from Baltic amber till now. The first two species of ladybird beetles from Eocene Baltic amber are described and placed in the genus Serangium, namely S. twardowskii sp. nov. and S. gedanicum sp. nov. Their phylogenetic placement in the subfamily Microweiseinae is provided. A key to the fossil species of Serangium is given. Extant representatives of the genus are distributed mainly in tropical areas of Asia and Oceania, and are specialised predators of whiteflies. Current discovery shows that during the Middle Eocene, the genus Serangium was distributed wider in the Northern Hemisphere and the evolution of these ladybird beetles was probably influenced by the evolution of whiteflies which are also found in the Baltic amber.