Arnold H. Staniczek

Coxoplectoptera, a new fossil order of Palaeoptera (Arthropoda: Insecta), with comments on the phylogeny of the stem group of mayflies (Ephemeroptera)

Mickoleitia longimanus gen. et sp.n. is described from the Lower Cretaceous limestone of the Crato Formation in Brazil. It is attributed to a new family Mickoleitiidae and a new fossil insect order Coxoplectoptera within the palaeopterous Ephemerida, based on the presence of an elongated costal brace. This fossil insect exhibits a very peculiar combination of derived characters like specialized forelegs with strongly elongated, free coxae, single-clawed pretarsus, and distinctly skewed pterothorax as in dragonflies. On the other hand, several plesiomorphies are present that exclude this taxon from modern Ephemeroptera, namely large hind wings with widened anal area and numerous cross veins that separate the elongate costal brace from the costal margin. Fossil larvae described by Willmann as larval Cretereismatidae are herein attributed to Mickoleitiidae fam.n., based on the shared presence of broad hind wing buds with distinctly broadened anal area, wing bud venation similar to the adult holotype, and subchelate forelegs with elongate free coxae. These larvae are also highly autapomorphic in the structure of their abdominal gills and laterally flattened body with vertically oval section that is unique within Ephemerida. On the other hand they possess plesiomorphic lateral wing pads with pronounced articulation like Palaeozoic pterygote larvae, while wing pads in modern insects are always secondarily fused to the tergum. A similar fossil larva from the Jurassic of Transbaikals was earlier described as Mesogenesia petersae and classified within modern mayflies. It is herein attributed to Mickoleitiidae fam.n. Coxoplectoptera are recognized as putative sister group of modern Ephemeroptera based on the shared presence of only 7 pairs of abdominal gills, while Permoplectoptera still have retained 9 pairs of gills. The phylogenetic reclassification of the mayfly stem group by Willmann is critically discussed and modified.

New fossil insect order Permopsocida elucidates major radiation and evolution of suction feeding in hemimetabolous insects (Hexapoda: Acercaria)

With nearly 100,000 species, the Acercaria (lice, plant lices, thrips, bugs) including number of economically important species is one of the most successful insect lineages. However, its phylogeny and evolution of mouthparts among other issues remain debatable. Here new methods of preparation permitted the comprehensive anatomical description of insect inclusions from mid-Cretaceous Burmese amber in astonishing detail. These “missing links” fossils, attributed to a new order Permopsocida, provide crucial evidence for reconstructing the phylogenetic relationships in the Acercaria, supporting its monophyly, and questioning the position of Psocodea as sister group of holometabolans in the most recent phylogenomic study. Permopsocida resolves as sister group of Thripida + Hemiptera and represents an evolutionary link documenting the transition from chewing to piercing mouthparts in relation to suction feeding. Identification of gut contents as angiosperm pollen documents an ecological role of Permopsocida as early pollen feeders with relatively unspecialized mouthparts. This group existed for 185 million years, but has never been diverse and was superseded by new pollenivorous pollinators during the Cretaceous co-evolution of insects and flowers. The key innovation of suction feeding with piercing mouthparts is identified as main event that triggered the huge post-Carboniferous radiation of hemipterans, and facilitated the spreading of pathogenic vectors.

Revision of the giant pterygote insect Bojophlebia prokopi Kukalová-Peck, 1985 (Hydropalaeoptera: Bojophlebiidae) from the Carboniferous of the Czech Republic, with the first cladistic analysis of fossil palaeopterous insects

The holotype is redescribed of the giant pterygote insect Bojophlebia prokopi Kukalová-Peck, 1985 from the Pennsylvanian of the Czech Republic. Multiple errors in the original description are documented and corrected. Bojophlebia prokopi 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 s.l. 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. http://zoobank.org/urn/urn:lsid:zoobank.org:pub:1B16D6AB-2B29-4891-A61D-7A9212A85493

Neither silverfish nor fowl: the enigmatic Carboniferous Carbotriplura kukalovae Kluge, 1996 (Insecta: Carbotriplurida) is the putative fossil sister group of winged insects (Insecta: Pterygota)

We revise the type material of the enigmatic fossil insect Carbotriplura kukalovae Kluge, 1996 from the Pennsylvanian of the Czech Republic. Multiple errors in the original description are documented and corrected. C. kukalovae is regarded as a possible transitional fossil link between Zygentoma and Pterygota. Carbotriplurida is therefore elevated to ordinal rank and considered as putative fossil sister group of Pterygota. The paranotal theory of the origin of insect wings and the parachute theory of origin of insect flight are briefly discussed and further corroborated. Testajapyx thomasi from the Pennsylvanian of Mazon Creek is tentatively considered as Dermaptera rather than Diplura.

Caenis vanuatensis, a new species of mayflies (Ephemeroptera: Caenidae) from Vanuatu

Abstract The first species of Caenidae from Espiritu Santo, Vanuatu, is described based on the larva and the female imago. Caenis vanuatensis sp. n. is most likely a parthenogenetic species, as only females have been found so far out of numerous specimens collected. The phylogenetic relationships of the new species thus remain enigmatic, as the distribution of Caenis throughout the Australian region is largely unknown and the taxonomically important males are missing.

Evolution of Baetiscidae (Ephemeroptera): current state of knowledge of the family

Recent discovery and description of fossil Baetiscidae has dramatically changed the biogeographical and evolutionary perspective of the family. Since the mid-eighteenth century until recently, the family has been known only from the Nearctic realm and all extant species of the family still remain geographically limited to this region. Here, we present the first parsimony algorithm generated cladistic analysis of the phylogeny of Baetisca based on external morphological characters of both larvae and adults. The results are compared with the previously published phylogeny of the genus and differences between the two studies are discussed. The southeastern United States is the centre of biodiversity of the genus where 11 of the 12 currently known nominal species have been reported. Biogeographical and evolutionary implications of the current geographic range of extant Baetisca and fossil species are discussed.

Baetis (Baetis) cypronyx sp. n., a new species of the Baetis alpinus species-group (Insecta, Ephemeroptera, Baetidae) from Cyprus, with annotated checklist of Baetidae in the Mediterranean islands

Abstract A detailed description of the larvae of Baetis (Baetis) cypronyx sp. n., a representative of the Baetis alpinus species-group within the mayfly family Baetidae, is provided, including a differential diagnosis with regard to closely related species of the group, especially Baetis melanonyx (Pictet, 1843) and Baetis baroukianus Thomas & Dia, 1984. The new species is mainly distinguished by mouthparts (i.e. the shape and setation of labrum, maxillary and labial palps, details of paraglossae and mandibular incisors), setation of legs and abdominal terga, and length of paracercus. All available data on the biology of this putative endemic species of Cyprus are summarized. Annotated distributional data of the 33 species of Baetidae so far recorded from the Mediterranean islands are given, including new records and also including first data from Malta.

Revision of fossil Metretopodidae (Insecta: Ephemeroptera) in Baltic amber – Part 2: Description of a new species of Metretopus Eaton, 1901

In the second part of the revision of fossil Metretopodidae, a new species of the genus Metretopus Eaton, 1901 is described and illustrated based on a male imago. Metretopus dividussp. nov. is the second fossil species of the genus. Distinguishing characters for its separation from other fossil and recent representatives of Metretopus are discussed. http://www.zoobank.org/urn:lsid:zoobank.org:act:BCC5CB69-8014-4DA1-A0DA-DCF714969081

Revision of fossil Metretopodidae (Insecta: Ephemeroptera) in Baltic amber-Part 3: Description of two new species of Siphloplecton Clemens, 1915, with notes on the re-discovered lectotype of Siphloplecton macrops (Pictet-Baraban & Hagen, 1856).

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.

Initial commented checklist of Iranian mayflies, with new area records and description of Procloeon caspicum sp. n. (Insecta, Ephemeroptera, Baetidae)

Abstract 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, sp. n. is described based on mature larva and egg. Critical differential diagnostic characters distinguishing the species from related taxa are discussed in detail.