The adult holotype of the fossil mayfly Astraeoptera cretacica Brandão et al. 2021 from the Creta... more The adult holotype of the fossil mayfly Astraeoptera cretacica Brandão et al. 2021 from the Cretaceous Crato Formation, Brazil, is reviewed and attributed to a new family Astraeopteridae fam. nov. Based on alate specimens, we also describe further new representatives of Astraeopteridae fam. nov., namely Astraeoptera vitrea sp. nov. and Astraeoptera oligovenata sp. nov., as well as the new genus and species Eosophobia acuta gen. et sp. nov. A subsequent character analysis of the new material suggests systematic affinities of Astraeopteridae fam. nov. with those extant families of Siphlonuroidea distributed in the Southern Hemisphere. These newly described fossil Siphlonuroidea from the Cretaceous of Brazil thus add to the biogeography and systematics of mayflies.
Among mayflies, the Hexagenitidae is the most common family of the Cretaceous, including in the w... more Among mayflies, the Hexagenitidae is the most common family of the Cretaceous, including in the world-renowned Crato Formation (Aptian) of northeastern Brazil. However, most of the Crato mayfly fau...
The adult holotype of the fossil mayfly Astraeoptera cretacica Brandão et al. 2021 from the Creta... more The adult holotype of the fossil mayfly Astraeoptera cretacica Brandão et al. 2021 from the Cretaceous Crato Formation, Brazil, is reviewed and attributed to a new family Astraeopteridae fam. nov. Based on alate specimens, we also describe further new representatives of Astraeopteridae fam. nov., namely Astraeoptera vitrea sp. nov. and Astraeoptera oligovenata sp. nov., as well as the new genus and species Eosophobia acuta gen. et sp. nov. A subsequent character analysis of the new material suggests systematic affinities of Astraeopteridae fam. nov. with those extant families of Siphlonuroidea distributed in the Southern Hemisphere. These newly described fossil Siphlonuroidea from the Cretaceous of Brazil thus add to the biogeography and systematics of mayflies.
Continental ecosystems of the middle Permian Period (273–259 million years ago) are poorly unders... more Continental ecosystems of the middle Permian Period (273–259 million years ago) are poorly understood. In South Africa, the vertebrate fossil record is well documented for this time interval, but the plants and insects are virtually unknown, and are rare globally. This scarcity of data has hampered studies of the evolution and diversification of life, and has precluded detailed reconstructions and analyses of ecosystems of this critical period in Earth’s history. Here we introduce a new locality in the southern Karoo Basin that is producing exceptionally well-preserved and abundant fossils of novel freshwater and terrestrial insects, arachnids, and plants. Within a robust regional geochronological, geological and biostratigraphic context, this Konservat- and Konzentrat-Lagerstätte offers a unique opportunity for the study and reconstruction of a southern Gondwanan deltaic ecosystem that thrived 266–268 million years ago, and will serve as a high-resolution ecological baseline toward...
The systematics of all known extant and fossil taxa of the mayfly family Baetiscidae is reviewed.... more The systematics of all known extant and fossil taxa of the mayfly family Baetiscidae is reviewed. Previously described fossil Baetiscidae are critically reviewed and their systematic position is re-evaluated. Added herein is the description of Balticobaetisca bispinatasp. nov., a new fossil mayfly from Eocene Baltic Amber. Also described for the first time is the putative adult of Protobaetisca bechlyi Staniczek, 2007 from the Cretaceous Crato Formation of Brazil. Based on external morphological characters of nymphs and adults, we present the first integrated cladistic analysis of both fossil and extant Baetiscidae, which revealed the monophyly of the respective fossil and extant genera and their phylogenetic splitting sequence as Cretomitarcys + (Protobaetisca + (Balticobaetisca + Baetisca)).
Proceedings of the Royal Society B: Biological Sciences, 2022
One key event in insect evolution was the development of mandibles with two joints, which allowed... more One key event in insect evolution was the development of mandibles with two joints, which allowed powerful biting but restricted their movement to a single degree of freedom. These mandibles define the Dicondylia, which constitute over 99% of all extant insect species. It was common doctrine that the dicondylic articulation of chewing mandibles remained unaltered for more than 400 million years. We report highly modified mandibles overcoming the restrictions of a single degree of freedom and hypothesize their major role in insect diversification. These mandibles are defining features of parasitoid chalcid wasps, one of the most species-rich lineages of insects. The shift from powerful chewing to precise cutting likely facilitated adaptations to parasitize hosts hidden in hard substrates, which pose challenges to the emerging wasps. We reveal a crucial step in insect evolution and highlight the importance of comprehensive studies even of putatively well-known systems.
<div><p>The holotype is redescribed of the giant pterygote insect <i>Bojophlebi... more <div><p>The holotype is redescribed of the giant pterygote insect <i>Bojophlebia prokopi</i> Kukalová-Peck, 1985 from the Pennsylvanian of the Czech Republic. Multiple errors in the original description are documented and corrected. <i>Bojophlebia prokopi</i> has neither any visible traces of a costal brace nor an anal brace, but it does show triadic branchings of MA, MP, CuA, and even, as rare a plesiomorphy, of CuP. It is therefore rejected as a fossil stem mayfly and attributed as sister group of all other Hydropalaeoptera. The first cladistic analysis of fossil palaeopterous insects, including different palaeodictyopterid groups, is presented. A revised phylogeny of Hydropalaeoptera and the stem line of Ephemeroptera are suggested. Palaeodictyopterida is recognized as sister group of Neoptera; thus Palaeoptera <i>s.l</i>. is rejected as a paraphyletic taxon. Four new higher taxa – Paranotalia, Euhydropalaeoptera, Neopterygota and Litophlebioidea superfam. nov. – are introduced, as well as the new family Lithoneuridae.</p><p><a href="http://zoobank.org/urn/urn:lsid:zoobank.org:pub:1B16D6AB-2B29-4891-A61D-7A9212A85493" target="_blank">http://zoobank.org/urn/urn:lsid:zoobank.org:pub:1B16D6AB-2B29-4891-A61D-7A9212A85493</a></p></div
An initial checklist of mayflies (Ephemeroptera) of Iran compiled based on the literature data an... more An initial checklist of mayflies (Ephemeroptera) of Iran compiled based on the literature data and new data from the north-western Iran reported only 46 species and 25 genera. Available literature data were often highly fragmented and referred to material limited to occasional collections with only several specimens examined. Moreover, many studies (mostly focusing on the water quality assessment based on macroinvertebrates) reported species or genera with distribution restricted to the Nearctic and Neotropic Region, suggesting that the authors used inappropriate determination keys. Most of valid records of mayflies were confined to the northern part of Iran, mostly Alborz Mts. To fill a significant gap in our knowledge on mayfly diversity, we started an extensive study covering the geographical and ecological gradients within Iran. Our field trips conducted in 2016–2018 set up a network of 234 localities to study the diversity and composition of mayfly assemblages in different type...
A new species of mayfly, Rallidens platydontis, from the South Island of New Zealand is described... more A new species of mayfly, Rallidens platydontis, from the South Island of New Zealand is described. The principal life stages are included and associated. Notes on ecology are given and a distribution map provided. Diagnostic characters of the new species are illustrated and compared with those of Rallidens mcfarlanei. Keys are provided for egg, larval and adult life stages.
In the aquatic insects, mayfly nymphs have developed a remarkable diversity of life strategies an... more In the aquatic insects, mayfly nymphs have developed a remarkable diversity of life strategies and feeding habits. However, the early evolution of these strategies and the role of mayfly nymphs in freshwater palaeoecosystems remain largely unknown. We investigated fossil mayfly nymphs from the Gr es a Voltzia Formation from the Middle Triassic, France, which represents the oldest known outcrop yielding a diverse mayfly fauna. We focused on three species, namely Mesoplectopteron longipes, Vogesonympha ludovici and Triassoephemera punctata, and investigated both original type material and numerous previously unstudied specimens. As a result, we provide a new interpretation of their morphology, life histories, and systematics: the nymphs of M. longipes, V. ludovici and T. punctata were aquatic and at least the former two species undoubtedly lived in flowing water as passive filter-feeders. This finding represents the so far earliest occurrence of this feeding strategy in aquatic insects. The most obvious morphological adaptations to this lifestyle are long filtering setae arranged in rows on forelegs, analogous to some modern mayfly nymphs. We place the three investigated species M. longipes, V. ludovici and T. punctata in a newly established, unranked higher taxon, Sinebranchia, representing the sister group to Heptabranchia (Coxoplectoptera + Ephemeroptera). Sinebranchia share a single-segmented tarsus synapomorphic with Heptabranchia, but retain the plesiomorphic condition of double tarsal claws. Sinebranchia are also characterized by their autapomorphic lack of tracheal gills, probably connected to their life in flowing, welloxygenated water.
The small, monophyletic mayfly family Vietnamellidae Allen, 1984 has so far only been known from ... more The small, monophyletic mayfly family Vietnamellidae Allen, 1984 has so far only been known from a few extant species of the genus Vietnamella Tshernova, 1972, which are all distributed in the Oriental Realm (Vietnam, Thailand, China, and India). Herein we report the first fossil record of Vietnamellidae based on a male and female imago from Mid-Cretaceous Burmese amber. We establish the new genus Burmellagen. nov. to accommodate these two new Mesozoic specimens. Their attribution to Vietnamellidae is supported by the rounded shape of the hind wings with arched outer margin, the course of thoracic sutures, and characteristics of venation, especially of MP and Cu of the forewings and associated intercalary veins of the cubital field. At the same time, Burmellagen. nov. clearly differs from Vietnamella by a diminished number of longitudinal and cross veins in the hind wings, and by the different shape of male genitalia. This first fossil record of Vietnamellidae supports an age of at ...
Emerging from the sixteenth century “Cabinet of Arts and Natural Curiosities” of the Dukes of Wur... more Emerging from the sixteenth century “Cabinet of Arts and Natural Curiosities” of the Dukes of Wurttemberg, the Stuttgart State Museum of Natural History (SMNS) ranks among the oldest and most prominent natural history museums in Europe. With its collections, libraries, laboratories, and other research facilities, the museum comprises two building complexes both with extensive exhibition and storage areas. The permanent exhibitions at Museum Schloss Rosenstein and Museum am Lowentor are tightly interwoven in form and content. Today, the SMNS collections hold more than 12 million specimens and other natural objects as well as numerous associated data on taxonomy, genetics, ecology, and geography. Together these comprehensive records and archives of life on Earth and its history constitute a large-scale research infrastructure used by research scientists and the international community. Characteristic of the SMNS is the tight collaboration between its paleontologists and researchers working on extant fauna.
Combining morphological and molecular data in an integrative approach, three new mayfly species o... more Combining morphological and molecular data in an integrative approach, three new mayfly species of Epeorus (Caucasiron) are described. These include Epeorus (Caucasiron) alborzicus Hrivniak & Sroka, sp. nov. and Epeorus (Caucasiron) shargi Hrivniak & Sroka, sp. nov. from northern Iran, and Epeorus (Caucasiron) zagrosicus Hrivniak & Sroka, sp. nov. from central Iran. They are unambiguously delimited using both distance-based and likelihood-based approaches in the analyses of barcode COI sequences. Each new species is compared with other species of the subgenus and morphological diagnostic characters are provided. Based on extensive sampling of streams throughout the country, the distribution and habitat preferences of all Caucasiron species in Iran are assessed. Altogether, there are now six species recorded, among them also E. (C.) nigripilosus Sinitshenkova, 1976 is reported for the first time in Iran. Five species are distributed in the Alborz Mts. in northern Iran, one species wa...
The adult holotype of the fossil mayfly Astraeoptera cretacica Brandão et al. 2021 from the Creta... more The adult holotype of the fossil mayfly Astraeoptera cretacica Brandão et al. 2021 from the Cretaceous Crato Formation, Brazil, is reviewed and attributed to a new family Astraeopteridae fam. nov. Based on alate specimens, we also describe further new representatives of Astraeopteridae fam. nov., namely Astraeoptera vitrea sp. nov. and Astraeoptera oligovenata sp. nov., as well as the new genus and species Eosophobia acuta gen. et sp. nov. A subsequent character analysis of the new material suggests systematic affinities of Astraeopteridae fam. nov. with those extant families of Siphlonuroidea distributed in the Southern Hemisphere. These newly described fossil Siphlonuroidea from the Cretaceous of Brazil thus add to the biogeography and systematics of mayflies.
Among mayflies, the Hexagenitidae is the most common family of the Cretaceous, including in the w... more Among mayflies, the Hexagenitidae is the most common family of the Cretaceous, including in the world-renowned Crato Formation (Aptian) of northeastern Brazil. However, most of the Crato mayfly fau...
The adult holotype of the fossil mayfly Astraeoptera cretacica Brandão et al. 2021 from the Creta... more The adult holotype of the fossil mayfly Astraeoptera cretacica Brandão et al. 2021 from the Cretaceous Crato Formation, Brazil, is reviewed and attributed to a new family Astraeopteridae fam. nov. Based on alate specimens, we also describe further new representatives of Astraeopteridae fam. nov., namely Astraeoptera vitrea sp. nov. and Astraeoptera oligovenata sp. nov., as well as the new genus and species Eosophobia acuta gen. et sp. nov. A subsequent character analysis of the new material suggests systematic affinities of Astraeopteridae fam. nov. with those extant families of Siphlonuroidea distributed in the Southern Hemisphere. These newly described fossil Siphlonuroidea from the Cretaceous of Brazil thus add to the biogeography and systematics of mayflies.
Continental ecosystems of the middle Permian Period (273–259 million years ago) are poorly unders... more Continental ecosystems of the middle Permian Period (273–259 million years ago) are poorly understood. In South Africa, the vertebrate fossil record is well documented for this time interval, but the plants and insects are virtually unknown, and are rare globally. This scarcity of data has hampered studies of the evolution and diversification of life, and has precluded detailed reconstructions and analyses of ecosystems of this critical period in Earth’s history. Here we introduce a new locality in the southern Karoo Basin that is producing exceptionally well-preserved and abundant fossils of novel freshwater and terrestrial insects, arachnids, and plants. Within a robust regional geochronological, geological and biostratigraphic context, this Konservat- and Konzentrat-Lagerstätte offers a unique opportunity for the study and reconstruction of a southern Gondwanan deltaic ecosystem that thrived 266–268 million years ago, and will serve as a high-resolution ecological baseline toward...
The systematics of all known extant and fossil taxa of the mayfly family Baetiscidae is reviewed.... more The systematics of all known extant and fossil taxa of the mayfly family Baetiscidae is reviewed. Previously described fossil Baetiscidae are critically reviewed and their systematic position is re-evaluated. Added herein is the description of Balticobaetisca bispinatasp. nov., a new fossil mayfly from Eocene Baltic Amber. Also described for the first time is the putative adult of Protobaetisca bechlyi Staniczek, 2007 from the Cretaceous Crato Formation of Brazil. Based on external morphological characters of nymphs and adults, we present the first integrated cladistic analysis of both fossil and extant Baetiscidae, which revealed the monophyly of the respective fossil and extant genera and their phylogenetic splitting sequence as Cretomitarcys + (Protobaetisca + (Balticobaetisca + Baetisca)).
Proceedings of the Royal Society B: Biological Sciences, 2022
One key event in insect evolution was the development of mandibles with two joints, which allowed... more One key event in insect evolution was the development of mandibles with two joints, which allowed powerful biting but restricted their movement to a single degree of freedom. These mandibles define the Dicondylia, which constitute over 99% of all extant insect species. It was common doctrine that the dicondylic articulation of chewing mandibles remained unaltered for more than 400 million years. We report highly modified mandibles overcoming the restrictions of a single degree of freedom and hypothesize their major role in insect diversification. These mandibles are defining features of parasitoid chalcid wasps, one of the most species-rich lineages of insects. The shift from powerful chewing to precise cutting likely facilitated adaptations to parasitize hosts hidden in hard substrates, which pose challenges to the emerging wasps. We reveal a crucial step in insect evolution and highlight the importance of comprehensive studies even of putatively well-known systems.
<div><p>The holotype is redescribed of the giant pterygote insect <i>Bojophlebi... more <div><p>The holotype is redescribed of the giant pterygote insect <i>Bojophlebia prokopi</i> Kukalová-Peck, 1985 from the Pennsylvanian of the Czech Republic. Multiple errors in the original description are documented and corrected. <i>Bojophlebia prokopi</i> has neither any visible traces of a costal brace nor an anal brace, but it does show triadic branchings of MA, MP, CuA, and even, as rare a plesiomorphy, of CuP. It is therefore rejected as a fossil stem mayfly and attributed as sister group of all other Hydropalaeoptera. The first cladistic analysis of fossil palaeopterous insects, including different palaeodictyopterid groups, is presented. A revised phylogeny of Hydropalaeoptera and the stem line of Ephemeroptera are suggested. Palaeodictyopterida is recognized as sister group of Neoptera; thus Palaeoptera <i>s.l</i>. is rejected as a paraphyletic taxon. Four new higher taxa – Paranotalia, Euhydropalaeoptera, Neopterygota and Litophlebioidea superfam. nov. – are introduced, as well as the new family Lithoneuridae.</p><p><a href="http://zoobank.org/urn/urn:lsid:zoobank.org:pub:1B16D6AB-2B29-4891-A61D-7A9212A85493" target="_blank">http://zoobank.org/urn/urn:lsid:zoobank.org:pub:1B16D6AB-2B29-4891-A61D-7A9212A85493</a></p></div
An initial checklist of mayflies (Ephemeroptera) of Iran compiled based on the literature data an... more An initial checklist of mayflies (Ephemeroptera) of Iran compiled based on the literature data and new data from the north-western Iran reported only 46 species and 25 genera. Available literature data were often highly fragmented and referred to material limited to occasional collections with only several specimens examined. Moreover, many studies (mostly focusing on the water quality assessment based on macroinvertebrates) reported species or genera with distribution restricted to the Nearctic and Neotropic Region, suggesting that the authors used inappropriate determination keys. Most of valid records of mayflies were confined to the northern part of Iran, mostly Alborz Mts. To fill a significant gap in our knowledge on mayfly diversity, we started an extensive study covering the geographical and ecological gradients within Iran. Our field trips conducted in 2016–2018 set up a network of 234 localities to study the diversity and composition of mayfly assemblages in different type...
A new species of mayfly, Rallidens platydontis, from the South Island of New Zealand is described... more A new species of mayfly, Rallidens platydontis, from the South Island of New Zealand is described. The principal life stages are included and associated. Notes on ecology are given and a distribution map provided. Diagnostic characters of the new species are illustrated and compared with those of Rallidens mcfarlanei. Keys are provided for egg, larval and adult life stages.
In the aquatic insects, mayfly nymphs have developed a remarkable diversity of life strategies an... more In the aquatic insects, mayfly nymphs have developed a remarkable diversity of life strategies and feeding habits. However, the early evolution of these strategies and the role of mayfly nymphs in freshwater palaeoecosystems remain largely unknown. We investigated fossil mayfly nymphs from the Gr es a Voltzia Formation from the Middle Triassic, France, which represents the oldest known outcrop yielding a diverse mayfly fauna. We focused on three species, namely Mesoplectopteron longipes, Vogesonympha ludovici and Triassoephemera punctata, and investigated both original type material and numerous previously unstudied specimens. As a result, we provide a new interpretation of their morphology, life histories, and systematics: the nymphs of M. longipes, V. ludovici and T. punctata were aquatic and at least the former two species undoubtedly lived in flowing water as passive filter-feeders. This finding represents the so far earliest occurrence of this feeding strategy in aquatic insects. The most obvious morphological adaptations to this lifestyle are long filtering setae arranged in rows on forelegs, analogous to some modern mayfly nymphs. We place the three investigated species M. longipes, V. ludovici and T. punctata in a newly established, unranked higher taxon, Sinebranchia, representing the sister group to Heptabranchia (Coxoplectoptera + Ephemeroptera). Sinebranchia share a single-segmented tarsus synapomorphic with Heptabranchia, but retain the plesiomorphic condition of double tarsal claws. Sinebranchia are also characterized by their autapomorphic lack of tracheal gills, probably connected to their life in flowing, welloxygenated water.
The small, monophyletic mayfly family Vietnamellidae Allen, 1984 has so far only been known from ... more The small, monophyletic mayfly family Vietnamellidae Allen, 1984 has so far only been known from a few extant species of the genus Vietnamella Tshernova, 1972, which are all distributed in the Oriental Realm (Vietnam, Thailand, China, and India). Herein we report the first fossil record of Vietnamellidae based on a male and female imago from Mid-Cretaceous Burmese amber. We establish the new genus Burmellagen. nov. to accommodate these two new Mesozoic specimens. Their attribution to Vietnamellidae is supported by the rounded shape of the hind wings with arched outer margin, the course of thoracic sutures, and characteristics of venation, especially of MP and Cu of the forewings and associated intercalary veins of the cubital field. At the same time, Burmellagen. nov. clearly differs from Vietnamella by a diminished number of longitudinal and cross veins in the hind wings, and by the different shape of male genitalia. This first fossil record of Vietnamellidae supports an age of at ...
Emerging from the sixteenth century “Cabinet of Arts and Natural Curiosities” of the Dukes of Wur... more Emerging from the sixteenth century “Cabinet of Arts and Natural Curiosities” of the Dukes of Wurttemberg, the Stuttgart State Museum of Natural History (SMNS) ranks among the oldest and most prominent natural history museums in Europe. With its collections, libraries, laboratories, and other research facilities, the museum comprises two building complexes both with extensive exhibition and storage areas. The permanent exhibitions at Museum Schloss Rosenstein and Museum am Lowentor are tightly interwoven in form and content. Today, the SMNS collections hold more than 12 million specimens and other natural objects as well as numerous associated data on taxonomy, genetics, ecology, and geography. Together these comprehensive records and archives of life on Earth and its history constitute a large-scale research infrastructure used by research scientists and the international community. Characteristic of the SMNS is the tight collaboration between its paleontologists and researchers working on extant fauna.
Combining morphological and molecular data in an integrative approach, three new mayfly species o... more Combining morphological and molecular data in an integrative approach, three new mayfly species of Epeorus (Caucasiron) are described. These include Epeorus (Caucasiron) alborzicus Hrivniak & Sroka, sp. nov. and Epeorus (Caucasiron) shargi Hrivniak & Sroka, sp. nov. from northern Iran, and Epeorus (Caucasiron) zagrosicus Hrivniak & Sroka, sp. nov. from central Iran. They are unambiguously delimited using both distance-based and likelihood-based approaches in the analyses of barcode COI sequences. Each new species is compared with other species of the subgenus and morphological diagnostic characters are provided. Based on extensive sampling of streams throughout the country, the distribution and habitat preferences of all Caucasiron species in Iran are assessed. Altogether, there are now six species recorded, among them also E. (C.) nigripilosus Sinitshenkova, 1976 is reported for the first time in Iran. Five species are distributed in the Alborz Mts. in northern Iran, one species wa...
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