Papers by Arnold H Staniczek
May ies (Ephemeroptera) are among the oldest pterygote insects, with the earliest fossils dating ... more May ies (Ephemeroptera) are among the oldest pterygote insects, with the earliest fossils dating back to the Late Carboniferous. Within may ies, Leptophlebiidae are a highly diverse and widespread group, with approximately 140 genera and 640 species. Whereas taxonomy, systematics, and phylogeny of extant Leptophlebiidae are in the focus of extensive studies, little is known about leptophlebiid fossil taxa. Because fossil remains of Ephemeroptera in sedimentary rocks are relatively rare, inclusions of may ies in amber are a unique source of information on their evolution and diversity in the past. Leptophlebiidae found in Cenozoic resins mostly belong to the subfamilies Leptophlebiinae (in Eocene Baltic amber) and Atalophlebiinae (in Miocene Dominican and Mexican ambers). In the present contribution, we con rm the rst nding of the genus Calliarcys from Eocene Baltic amber by using Micro-CT, which allowed con rming its generic placement by visualizing diagnostic key characters otherwise hidden by a cloud of turbidity. The description and morphological reconstruction of the new fossil species Calliarcys antiquus sp. nov. and new molecular data of the extant Calliarcys humilis Eaton, 1881 from the Iberian Peninsula do not support a distinct subfamily Calliarcyinae, but reveal close relationships to the genera Leptophlebia and Paraleptophlebia, thus suggesting a phylogenetic position of Calliarcys within Leptophlebiinae.
FIG. 4. — A, D, G, J, Vanuatubasis bidens (Kimmins, 1958) n. comb., holotype, ♂ (BMNH), anal appe... more FIG. 4. — A, D, G, J, Vanuatubasis bidens (Kimmins, 1958) n. comb., holotype, ♂ (BMNH), anal appendages in dorsal and lateral view; B, E, H, K, V. malekulana (Kimmins, 1936) n. comb., holotype, ♂ (BMNH), anal appendages in dorsal and lateral view; C, F, I, L, V. santoensis n. gen., n. sp., holotype, ♂ (SMNS: ODO 000242 K), anal appendages in dorsal and lateral views. Scale bars: 0.2 mm.
Integrative Systematics: Stuttgart Contributions to Natural History
Mitteilungen des Entomologischen Vereins Stuttgart, 2023
The Mantid fauna of Europe is rapidly changing due to the climate change of the last decades. Not... more The Mantid fauna of Europe is rapidly changing due to the climate change of the last decades. Not only Mantis religiosa, our single native species in Germany, is dynamically expanding, also Mediter-ranean and Asiatic species are extending their distributional range. Here, the first record of the Asian Giant Mantis, Hierodula tenuidentata Saussure, 1869, is reported for Germany.
Scientific Reports, Jul 20, 2023
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. The Brazilian Araripe Basin with its Crato Formation represents one of the most important Lagerstätte for Cretaceous insects 1,2. Particularly mayflies (Insecta, Ephemeroptera) are found most abundantly in these sedimentary limestones 3,4. The dominant autochthonous part of the Crato mayfly fauna is represented by the species Protoligoneuria limai Demoulin, 1955 of the extinct family Hexagenitidae Lameere, 1917 5. Nonetheless, nymphal and adult life stages of several other allochthonous groups can also be found. It is assumed that the nymphs were washed out from distantly connected lotic water bodies, while alate adults were reaching the depositional site through active flight 6,7. Among the allochthonous mayfly groups preserved in the limestones of the Crato Formation are Baetiscidae
Annales Zoologici, Mar 1, 2017
The Neotropical mayfly genus Borinquena Traver, 1938 so far comprises three described extant spec... more The Neotropical mayfly genus Borinquena Traver, 1938 so far comprises three described extant species from the Caribbean basin. Furthermore, three fossil species from Dominican amber have been described in this genus. Based on a single male imago, Borinquena schawallfussi sp. nov. is described in this contribution as the fourth fossil species of Borinquena from Miocene Dominican amber. The new species clearly differs from all other fossil and recent representatives of the genus Borinquena in forewing colouration and venation, hind wing shape and venation, and in the shape of penis lobes and their ventral spines.
Research Square (Research Square), Apr 7, 2022
May ies (Ephemeroptera) are among the oldest pterygote insects, with the earliest fossils dating ... more May ies (Ephemeroptera) are among the oldest pterygote insects, with the earliest fossils dating back to the Late Carboniferous. Within may ies, Leptophlebiidae are a highly diverse and widespread group, with approximately 140 genera and 640 species. Whereas taxonomy, systematics, and phylogeny of extant Leptophlebiidae are in the focus of extensive studies, little is known about leptophlebiid fossil taxa. Because fossil remains of Ephemeroptera in sedimentary rocks are relatively rare, inclusions of may ies in amber are a unique source of information on their evolution and diversity in the past. Leptophlebiidae found in Cenozoic resins mostly belong to the subfamilies Leptophlebiinae (in Eocene Baltic amber) and Atalophlebiinae (in Miocene Dominican and Mexican ambers). In the present contribution, we con rm the rst nding of the genus Calliarcys from Eocene Baltic amber by using Micro-CT, which allowed con rming its generic placement by visualizing diagnostic key characters otherwise hidden by a cloud of turbidity. The description and morphological reconstruction of the new fossil species Calliarcys antiquus sp. nov. and new molecular data of the extant Calliarcys humilis Eaton, 1881 from the Iberian Peninsula do not support a distinct subfamily Calliarcyinae, but reveal close relationships to the genera Leptophlebia and Paraleptophlebia, thus suggesting a phylogenetic position of Calliarcys within Leptophlebiinae.
Zootaxa, Apr 11, 2016
In this contribution, we provide a complementary description of the re-discovered lectotype of Si... more In this contribution, we provide a complementary description of the re-discovered lectotype of Siphloplecton macrops (Pictet-Baraban & Hagen, 1856). Additionally, complementary descriptions of S. barabani Staniczek & Godunko, 2012 and S. picteti Staniczek & Godunko, 2012 based on new material are given. Two new species, S. sartorii sp. nov. and S. gattolliati sp. nov., are described from male imagines and attributed to two newly defined species groups within Siphloplecton.
Scientific Reports
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.
ZooKeys, 2018
An initial checklist of mayflies (Ephemeroptera) of Iran is compiled based on critical review of ... more An initial checklist of mayflies (Ephemeroptera) of Iran is compiled based on critical review of available literature data, complemented with new data from 38 localities of Gilan and Ardabil provinces. At present, altogether only 46 species and 25 genera are known from Iran, 18 species are reported as new to Iran in this study. Some previously published data are critically evaluated and doubtful taxa are excluded from the list. Basic analysis of the distribution and biogeography of recorded species is given. Procloeon (Pseudocentroptilum) caspicum Sroka, is described based on mature larva and egg. Critical differential diagnostic characters distinguishing the species from related taxa are discussed in detail.
Zoologischer Anzeiger, 2000
The mandibles of several Ephemeroptera larvae have been investigated in order to reconstruct the ... more The mandibles of several Ephemeroptera larvae have been investigated in order to reconstruct the groundplan of the mandibular muscles and of the mandibular articulation of Ephemeroptera. The results indicate the presence of three points of mandibular ...
Integrative systematics, May 29, 2023
The function of wing bullae in mayflies (Insecta: Ephemeroptera) reveals new insights into the early evolution of Pterygota, 2023
Mayflies are basal winged insects of crucial importance for the understanding of the early evolut... more Mayflies are basal winged insects of crucial importance for the understanding of the early evolution of Pterygota. Unlike all other insects, they have two successive winged stages, the subimago and the imago. Their forewings feature so-called bullae, which are desclerotized spots in the anterior main veins. Up to now, they have been considered to play a major role in wing bending during flight.
We investigated bullae by multiple methods to reveal their structure and arrangement and to gain new
information on the evolution of insect flight. Bullae are mostly present in the anterior negative wing veins, disrupting
the otherwise rigid veins. High-speed videography reveals that mayfly wings do not bend during flight. Likewise,
different arrangements of bullae in different species do not correlate with different modes of flying. Observations
on the moulting of subimagines unravel that they are essential for wing bending during the extraction of the imaginal
wing from the subimaginal cuticle. Bullae define predetermined bending lines, which, together with a highly
flexible wing membrane enriched with resilin, permit wing bending during subimaginal moulting. Bullae are
only absent in those species that remain in the subimaginal stage or that use modified modes of moulting. Bullae are
also visible in fossil mayflies and can be traced back to stemgroup mayflies of the Early Permian, the 270 million years
old Protereismatidae, which most probably had bullae in both fore- and hind wings.
Bullae in mayfly wings do not play a role in flight as previously thought, but are crucial for wing bending
during subimaginal moulting. Thus, the presence of bullae is a reliable morphological marker for a subimaginal life
stage, confirming the existence of the subimago already in Permian Protereismatidae. A thorough search for bullae
in fossils of other pterygote lineages may reveal wheather they also had subimagines and at what point in evolution
this life stage was lost. In mayflies, however, the subimago may have been retained due to selective advantages
in connection with the transition from aquatic to terrestrial life or due to morphological requirements for a specialized
mating flight.
Integrative Systematics: Stuttgart Contributions to Natural History, 2021
BioOne Complete (complete.BioOne.org) is a full-text database of 200 subscribed and open-access t... more BioOne Complete (complete.BioOne.org) is a full-text database of 200 subscribed and open-access titles in the biological, ecological, and environmental sciences published by nonprofit societies, associations, museums, institutions, and presses.
Communications Biology
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...
Scientific Reports, Sep 8, 2022
Mayflies (Ephemeroptera) are among the oldest pterygote insects, with the earliest fossils dating... more Mayflies (Ephemeroptera) are among the oldest pterygote insects, with the earliest fossils dating back to the Late Carboniferous. Within mayflies, Leptophlebiidae are a highly diverse and widespread group, with approximately 140 genera and 640 species. Whereas taxonomy, systematics, and phylogeny of extant Leptophlebiidae are in the focus of extensive studies, little is known about leptophlebiid fossil taxa. Because fossil remains of Ephemeroptera in sedimentary rocks are relatively rare, inclusions of mayflies in amber are a unique source of information on their evolution and diversity in the past. Leptophlebiidae found in Cenozoic resins mostly belong to the subfamilies Leptophlebiinae (in Eocene Baltic amber) and Atalophlebiinae (in Miocene Dominican and Mexican ambers). In the present contribution, we confirm the first finding of the genus Calliarcys from Eocene Baltic amber by using Micro-CT, which allowed confirming its generic placement by visualizing diagnostic key characters otherwise hidden by a cloud of turbidity. Additionally, we present first molecular data on the extant species Calliarcys humilis Eaton, 1881 from the Iberian Peninsula and the barcode gap analysis for Leptophlebiinae and Habrophlebiinae. The genus Calliarcys Eaton, 1881 was established for the species C. humilis Eaton, 1881, which was described based on male and female adults from Spain 1. A detailed overview on taxonomy, distribution, and current knowledge of C. humilis was given by Godunko et al. 2. Originally thought to be an isolated West Palearctic species, more than 130 years later a second extant species, Calliarcys van Godunko and Bauernfeind, 2015, was described from two quite isolated localities in Turkey, namely in the Eastern Mediterranean (Izmir Province; W Turkey) and in East Anatolia (Bitlis Province). Its description led to a revision of diagnostic characters for Calliarcys and to the conclusion that Calliarcys has rather circum-Mediterranean origins 2. However, since the original description of the genus, its phylogenetic position within Leptophlebiidae remained enigmatic and controversial. The higher classification of Leptophlebiidae proposed by Peters 3 recognized only two subfamilies, Leptophlebiinae and Atalophlebiinae. Discussing the systematic position of some extinct Mesozoic and Cenozoic taxa, Kluge 4 only recognized Leptophlebiinae as monophyletic group and considered Atalophlebiinae as paraphyletic. Later, Kluge suggested a split of Leptophlebiinae s.l. into two subfamilies, Habrophlebiinae and Leptophlebiinae s.str. 4. Peters and Gilles 5 introduced the presence of squared ommatidia of the upper portion of the male compound eyes as convincing autapomorphic character supporting the monophyly of Atalophlebiinae.
Papers in Palaeontology
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.
Der Larvenkopf der neuseeländischen Eintagsfliege Oniscigaster wakefieldi wird beschrieben. Davon... more Der Larvenkopf der neuseeländischen Eintagsfliege Oniscigaster wakefieldi wird beschrieben. Davon ausgehend wird der Versuch unternommen, im Sinne der phylogenetischen Systematik Aussagen über den Bau des Larvenkopfes im Grundplan der Ephemeroptera zu treffen. Besonderes Gewicht liegt dabei auf der Anatomie und funktionellen Morphologie der larvalen Mundwerkzeuge und der Homologisierung ihrer Muskulatur. Im Mittelpunkt steht hierbei die Mandibel, an der ein vorderer Artikulationskomplex aus zwei Gelenkpunkten nachgewiesen wird. Zur Bewertung der gewonnenen Ergebnisse werden diese mit der Mandibel des Silberfischchens Tricholepidion gertschi und anderer Zygentoma verglichen. Alle Zygentoma weisen an der Mandibel ebenfalls einen anterioren Artikulationskomplex auf. Diese Übereinstimmungen werden als Grundplanmerkmal der Dicondylia gewertet. Darauf beruhend wird die Merkmalstransformation von der monocondylen Mandibel der Ectognatha bis zur dicondylen Mandibel der Odonata und Neoptera dargelegt und in ihrem funktionellen und phylogenetischen Kontext interpretiert. Dieser Umbau der Mandibel und ihrer Muskulatur ist korreliert mit Veränderungen im Bau und der Lage der vorderen Tentorialarme und der Kopfseitenwand. Keine der untersuchten Mandibeln ist segmentiert. Die Hypothese einer primär dreigliedrigen, telognathen Mandibel im Grundplan der Insekten wird daher zurückgewiesen. Die verschmolzenen Laden der larvalen Eintagsfliegenmaxille werden mit denen der Libellen verglichen und unterschiedliche Interpretationsmöglichkeiten der Befunde aufgezeigt. Die Ergebnisse dieser Untersuchung werden zusammen mit allen bisher diskutierten Merkmalen für eine phylogenetische Analyse der umstrittenen Verwandtschaftsverhältnisse zwischen den basalen Gruppen der Pterygota herangezogen. Aus diesen Daten ergibt sich als schlüssigste Verwandtschaftshypothese ein Schwestergruppenverhältnis zwischen den Ephemeroptera und allen anderen geflügelten Insekten, den Metapterygota.The larval head of the New Zealand mayfly Oniscigaster wakefieldi is investigated, a mayfly which is thought of as plesiomorphic in many respects. Based on a broad comparison with other mayfly species, it is aimed to reconstruct the groundplan of the larval head of mayflies. Emphasis is laid on the anatomy and functional morphology of the larval mouthparts and the homologization of their muscles. The results indicate the presence of an anterior articulation complex of the mandible that is made up of two points of mandibular attachment to the cranium in the groundplan of Ephemeroptera. These attachment points are compared with the condition in the relic silverfish Tricholepidion gertschi and other Zygentoma. It is shown that in the Zygentoma there is a similar articulation complex present. This correspondence is regarded as a groundplan character of Dicondylia. The character transformation from the monocondylous mandibles of Ectognatha to the dicondylous mandible of Odonata and Neoptera is interpreted in its functional and phylogenetic context. The alterations of the mandible and its muscles are correlated with changes in the construction and position of the anterior tentorial arms and the lateral cranium. None of the investigated mandibles show traces of segmentation. Thus the hypothesis of a tripartite telognathic mandible in the groundplan of insects is rejected. The fused galeolacinia of mayfly larvae is compared to the ones of the Odonata. The different ways of interpretation of these results are discussed. The findings of this study, together with all previously discussed characters, are used for a phylogenetic analysis to unravel the phylogenetic affinities between the three basal groups of the Pterygota (Ephemeroptera, Odonata, Neoptera). The data point to a sistergroup relationship between the Ephemeroptera and all other pterygote insects, the Metapterygota
Zoological Collections of Germany, 2018
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.
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Papers by Arnold H Staniczek
We investigated bullae by multiple methods to reveal their structure and arrangement and to gain new
information on the evolution of insect flight. Bullae are mostly present in the anterior negative wing veins, disrupting
the otherwise rigid veins. High-speed videography reveals that mayfly wings do not bend during flight. Likewise,
different arrangements of bullae in different species do not correlate with different modes of flying. Observations
on the moulting of subimagines unravel that they are essential for wing bending during the extraction of the imaginal
wing from the subimaginal cuticle. Bullae define predetermined bending lines, which, together with a highly
flexible wing membrane enriched with resilin, permit wing bending during subimaginal moulting. Bullae are
only absent in those species that remain in the subimaginal stage or that use modified modes of moulting. Bullae are
also visible in fossil mayflies and can be traced back to stemgroup mayflies of the Early Permian, the 270 million years
old Protereismatidae, which most probably had bullae in both fore- and hind wings.
Bullae in mayfly wings do not play a role in flight as previously thought, but are crucial for wing bending
during subimaginal moulting. Thus, the presence of bullae is a reliable morphological marker for a subimaginal life
stage, confirming the existence of the subimago already in Permian Protereismatidae. A thorough search for bullae
in fossils of other pterygote lineages may reveal wheather they also had subimagines and at what point in evolution
this life stage was lost. In mayflies, however, the subimago may have been retained due to selective advantages
in connection with the transition from aquatic to terrestrial life or due to morphological requirements for a specialized
mating flight.
We investigated bullae by multiple methods to reveal their structure and arrangement and to gain new
information on the evolution of insect flight. Bullae are mostly present in the anterior negative wing veins, disrupting
the otherwise rigid veins. High-speed videography reveals that mayfly wings do not bend during flight. Likewise,
different arrangements of bullae in different species do not correlate with different modes of flying. Observations
on the moulting of subimagines unravel that they are essential for wing bending during the extraction of the imaginal
wing from the subimaginal cuticle. Bullae define predetermined bending lines, which, together with a highly
flexible wing membrane enriched with resilin, permit wing bending during subimaginal moulting. Bullae are
only absent in those species that remain in the subimaginal stage or that use modified modes of moulting. Bullae are
also visible in fossil mayflies and can be traced back to stemgroup mayflies of the Early Permian, the 270 million years
old Protereismatidae, which most probably had bullae in both fore- and hind wings.
Bullae in mayfly wings do not play a role in flight as previously thought, but are crucial for wing bending
during subimaginal moulting. Thus, the presence of bullae is a reliable morphological marker for a subimaginal life
stage, confirming the existence of the subimago already in Permian Protereismatidae. A thorough search for bullae
in fossils of other pterygote lineages may reveal wheather they also had subimagines and at what point in evolution
this life stage was lost. In mayflies, however, the subimago may have been retained due to selective advantages
in connection with the transition from aquatic to terrestrial life or due to morphological requirements for a specialized
mating flight.