The earliest known fossil beetle †
Coleopsis archaica
is re-examined using Reflectance Transformation Imaging (RTI). The morphological observations are evaluated with respect to phylogenetic ...implications and the early evolution of Coleoptera. †
Coleopsis archaica
belongs to an early Permian branch of beetles, outside a monophyletic unit comprising Coleoptera (in the widest sense) excluding †Tshekardocoleidae. This clade is mainly characterized by a complex of apomorphic features: elytra with epipleura and with a close fit with the posterior body, thus forming a tightly sealed subelytral space. In contrast to this, the elytra of †
C. archaica
and †Tshekardocoleidae cover the metathorax and abdomen in a loose tent-like manner and posteriorly distinctly surpass the abdominal apex. So far, no synapomorphies of the two taxa from the first half of the Permian have been identified. The very short and transverse pronotum is likely an autapomorphy of †
C. archaica
. A thorough documentation of the structural features of early beetle fossils should have high priority. RTI is a very promising tool to obtain new and well-founded morphological data, which will allow a thorough phylogenetic evaluation of Permian beetles in future studies. We extended the conventional RTI workflow by focus merging and panoramic stitching, in order to overcome previous limitations. Taxonomic re-arrangements of stem group beetles including †
C. archaica
were suggested in recent studies by A.G. Kirejtshuk and co-workers. As they are not based on shared derived features they are irrelevant in a phylogenetic and evolutionary context.
The fossil record of dragonfly relatives (Odonatoptera) dates back to the Carboniferous, yet knowledge about these extinct animals is meagre. For most of the species little is known except for the ...characteristics of the wing venation. As a result, it is difficult to include fossil larvae in a (wing character based) phylogenetic tree as the wing venation is not visible in most of the larval instars. Two larval specimens from Cretaceous Burmese amber are in the focus of this study. The two specimens likely represent two subsequent early stage larval instars of the same individual. Not only is this an exceptional case to study ontogenetic processes in fossils – the larval instars are morphologically completely different from all known larvae of Odonata with respect to the posterior abdominal region. Therefore, besides the difficulties regarding the phylogenetic interpretation and though all Burmese amber odonatans are known from adults only, a new species – Arcanodraco filicauda n. sp. – is formally described. Aside from likely representing a new species, the morphology of the posterior abdominal region is highly informative for reconstructing the character evolution within the lineage towards modern dragonflies and damselflies. A long median process in both of the fossils meets all criteria to be interpreted as a terminal filum (structure or derivative of tergite of abdominal segment 11, annulated in one of the specimens). Although the exact phylogenetic affinity of Arcanodraco filicauda n. sp. remains enigmatic, the presence of a larval terminal filum can be reconstructed for the ground pattern of Odonatoptera (including its direct stem lineage).
Abstract Two fossils from Burmese amber are the subject of this study. The specimens differ in size; yet, they appear to be conspecific because of the profound morphological similarity. The fossils ...are interpreted as representatives of Isopoda, more precisely of the group Cymothoida, due to the presence of a triangular basipod of the uropod. Cymothoida comprises parasitic forms of Isopoda as well as many other types of feeding-habits. The morphology in the studied fossils suggests that they are not representatives of any of the parasitic ingroups of Cymothoida. Since there are no other findings of Isopoda from the Cretaceous with the same morphological features, the fossils at hand are described as a new species - Electrolana madelineae sp. nov. The smaller specimen lacks well-developed walking appendages on trunk segment seven; it can thus be interpreted as a manca stage (immature) individual. The systematic affinity and the functional morphology of the herein described fossils, as well as three seed shrimps (Ostracoda) in close proximity to one of the specimens, and the presence of pyrite in the amber piece points towards an aquatic lifestyle and a preservation in moist conditions. In addition, we review the fossil record of immature forms of Isopoda.
Within Neuroptera, the group of lacewings, the ingroup Nevrorthidae is special in several aspects: 1) the group may represent the sister group to all other neuropterans; 2) the group is quite ...species-poor with only 19 extant species known so far; 3) its representatives show a disjunct relict distribution; 4) it has very unusual appearing larvae. These larvae are very elongate, almost worm-shaped. Our overall knowledge of these larval forms is still very limited. We here review all known occurrences of extant larvae, amended by fossil larvae, all preserved in amber. So far there have been only 17 extant larval specimens of Nevrorthidae depicted in the literature. We also present new fossil larvae, with this reaching a total number of 16 known fossil larvae of Nevrorthidae. When plotting measured lengths, larvae cluster into more than three clusters. Also reconstructed size gains between these clusters indicate that Nevrorthidae might in fact develop through more than three larval stages, unlike other lacewings. A special subdivision of the trunk segments observed in many larvae indicates a specialisation for moving in confined spaces. Comparison of syn-inclusions and literature data make it likely that not all larvae lived in running waters, as previously assumed. Overall our knowledge remains rather limited, yet the data allow to draw some new conclusions about the life and evolution of these larval forms.
A collection of exceptionally well-preserved fossil specimens of crustaceans, clearly representatives of Isopoda, is presented here. Excavated from the late Eocene (approximately 40 million years ...ago) freshwater sediments of the Trupelník hill field site near Kučlín, Czech Republic, these specimens are preserved with many details of the appendages. The morphological characteristics of the fossils were documented using macro-photography with polarised light, as well as stereo imaging. These characteristics, especially including the trunk appendage morphology, were compared to those of related extant groups from different ontogenetic stages. All specimens are conspecific, representing a single species
gen. et sp. nov. Morphometric analysis of body shapes and sizes of the reconstructed fossils and related extant species were performed. These analyses provided insight into the ontogenetic stages of each reconstructed fossil specimen. In combination with the morphological assessment, the results indicate that the fossils represent at least two (possibly three) developmental stages, including immatures. The morphology of the appendages suggests that these fossils were parasites. The fossils are interpreted as either representatives of Cymothoidae or at least closely related to this group.
Two new species of fossil dragonflies from the Middle Miocene fossil site Schaßbach (Carinthia, Austria) are described. The presence of Gomphaeschna carinthiae sp. nov. and Ictinogomphus hassleri sp. ...nov. in the fossil record of Central Europe confirms the scenario of a more widespread distribution of the represented genera in the Miocene in contrast to their Recent distribution.
Burmese Cretaceous amber (~99 Ma, Myanmar) is famous for the preservation of a wide range of fauna and flora, including representatives of marine, freshwater and terrestrial groups. Here, we report ...on three ostracod specimens, that came visible as syninclusions to an aquatic isopod. The three specimens represent three different taxa, that were found preserved in a single piece of amber. One of the described specimens was studied using microCT scanning data. On the basis of general carapace morphology we assign all three to the group Podocopida, and (tentatively) its ingroup Cypridocopina. A lack of visibility of more particular diagnostic features such as adductor muscle scars and details of the marginal zone precludes a further identification, but we discuss possible affinities with either the marine-brackish group Pontocypridoidea or the non-marine group Cypridoidea. The taphonomy indicates that the studied ostracods had been subject to limited (if any) post-mortem transport, which could be consistent with marginal marine environments.
Larvae of flies and gnats (Diptera) form a crucial component of many terrestrial and freshwater ecosystems in the extant biosphere. Larvae of Diptera play a central role in water purification, matter ...and energy transfer in riparian ecosystems in rivers, carbon cycling in lakes and forests as well as being major decomposers of dead organic matter. Despite all these important roles, dipteran larvae are most often ignored in palaeoecological studies, due to the difficulty of the taxonomic identification of fossil larvae, but also due to the perceived importance of adult dipterans in palaeoentomological and taxonomic studies. Despite that, much information on palaeoecosystems can be gained from studying fossil dipteran larvae, in particular for well preserved specimens from fossil resins (ambers and copals). Since ambers are selectively preserving fauna of trunks and leaf litter, it allows us to learn a lot about xylophages and saprophages of amber forests, such as Eocene Baltic amber forest. Here we present immature stages (larvae and pupae) of the dipteran ingroup Bibionomorpha, from Baltic and Bitterfeld amber forests. We have recorded at least four different larval morphotypes, one with four distinct instars, and at least three pupal morphotypes. One larva is recognised as a new species and can be interpreted either as a representative of a highly derived ingroup of Bibionidae or as a sister species to Bibionidae. Also represented by single larval specimens are the groups Pachyneura (Pachyneuridae) and Sylvicola (Anisopodidae). The majority of the recorded specimens are representatives of the group Mycetobia (Anisopodidae). Due to the abundance of immature stages of Mycetobia, we have been able to reconstruct the number of larval stages (4) and relative growth rate of these fossil dipterans. We discuss implications of these finds.
The fossil record of Isopoda includes remains of presumed parasites. Among the fossils which have been discussed as potential parasites are those termed as Urda Münster, 1840. Some of these fossils ...have been discussed as possibly related to an extant group of parasites, Gnathiidae Leach, 1814. The type species of Urda – Urda rostrata Münster, 1840 – is herein interpreted as a close relative of the group Gnathiidae, based on the shared occurrence of a number of apomorphic features. This is with Urda punctata (Münster, 1842) herein being interpreted as a junior subjective synonym of U. rostrata. However, not all of the fossils associated with the name Urda can safely be identified as close relatives of Gnathiidae. Moreover, it is unclear whether the extinct species, which can be identified as close relatives of U. rostrata and Gnathiidae form a monophyletic group, as we could not identify an autapomorphy for a natural group Urda. A new species of close relatives of Urda rostrata and Gnathiidae – Urda buechneri n. sp. – is formally described based on µCT image data. Palaega suevica Reiff, 1936 and Palaega kessleri Reiff, 1936 are found to be subjective synonyms and are re-interpreted as Urda suevica n. comb. – a species closely related to U. rostrata. Due to the documented destruction of the holotype, a herein figured fossil specimen is designated as the neotype of Urda suevica. Palaega? stemmerbergensis Malzahn, 1968 is also interpreted as a close representative of U. rostrata and herein treated as Urda stemmerbergensis n. comb. Another already formally described species – Eobooralana rhodanica gen. et comb. nov. – is interpreted as a more distant relative, which is likely to be closer related to other extant species of Isopoda than those within Gnathiidae. For three species there are not enough characters preserved to interpret them as closely related to U. rostrata and Gnathiidae: Urda? liasica Frentzen, 1937 nom. dub. (type material destroyed, description insufficient for proper diagnosis), Urda? moravica Remeš, 1912 and Urda? zelandica Buckeridge and Johns, 1996.
Holometabola is a hyperdiverse group characterised by a strong morphological differentiation between early post-embryonic stages (= larvae) and adults. Adult forms of Holometabola, such as wasps, ...bees, beetles, butterflies, mosquitoes or flies, are strongly differentiated concerning their mouth parts. The larvae most often seem to retain rather plesiomorphic-appearing cutting-grinding mouth parts. Here we report a new unusual larva preserved in Burmese amber. Its mouth parts appear beak-like, forming a distinct piercing mouth cone. Such a morphology is extremely rare among larval forms, restricted to those of some beetles and lacewings. The mouth parts of the new fossil are forward oriented (prognathous). Additionally, the larva has distinct subdivisions of tergites and sternites into several sclerites. Also, the abdomen segments bear prominent protrusions. We discuss this unusual combination of characters in comparison to the many different types of holometabolan larvae. The here reported larva is a new addition to the 'unusual zoo' of the Cretaceous fauna including numerous, very unusual appearing forms that have gone extinct at the Cretaceous-Palaeogene boundary.
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