DOI: 10.3897/asp.84.e164719

Authors: Feng-E Li, Lin Yang, Jian-Kun Long, Zhi-Min Chang, Xiang-Sheng Chen

Abstract: Abstract The treehopper genus Tricentrus is a widespread group of insects, characterized by typical spines and, in some species, sexually dimorphic suprahumeral horns. With 238 described species worldwide, Tricentrus represents the most species-rich genus in the subfamily Centrotinae. Currently, 72 species are documented in China, yet the phylogenetic relationships and evolutionary history of this genus remain unknown. Here, we use an integrative approach to characterize and describe these treehoppers based on materials collected from China. We name three new species: Tricentrus allochrous Li & Chen sp. nov. and Tricentrus pianmaensis Li & Chen sp. nov. from Yunnan, Tricentrus dexingensis Li & Chen sp. nov. from Xizang, with the revision of one synonym. Combined analysis morphological comparisons, species delimitation and phylogeny reliably separate these new species from known species, and sexual dimorphism with suprahumeral horns in three species is reported for the first time. Divergence time estimation indicates that the Tricentrus differentiated during the Early Cretaceous (110.41 Mya). Ancestral state reconstruction reveals that the most recent common ancestor possessed suprahumeral horns in both sexes and exhibited a short tubular pygofer apex. Our analyses demonstrate four evolutionary transformations in pygofer apex morphology and seven distinct transitions in suprahumeral horn development. These morphological diversifications likely resulted from multiple mechanisms, including sexual selection, life history strategies with associated behavioral adaptations, and incomplete lineage sorting.

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DOI: 10.3897/asp.84.e154913

Authors: Gabriel S. Ferreira, André Silva Roza, José Ricardo M. Mermudes

Abstract: Abstract Coccoderus Buquet, 1840 is one of the largest genera in Torneutini (Cerambycinae), composed of 14 species. A previous morphology-based phylogeny recovered the genus as a monophyletic group. Since then, four species were described in the genus. Herein, we expanded this previous morphological dataset with the inclusion of seven characters and eight species. We performed maximum parsimony, maximum likelihood and Bayesian inference analyses. We used the Statistical Dispersal-Vicariance Analysis and the Bayesian Binary Method to reconstruct the potential ancestral ranges of Coccoderus. Our phylogenetic analysis recovered Coccoderus as a monophyletic group and shows shifts in species relationships, with two clades presenting new placements compared to the previous analysis. Biogeographic analyses identified the Chacoan dominion as a key region in the diversification of the genus, representing its ancestral area. Also, we described Coccoderus trimaculatus sp. nov., from Brazil, increasing the known species of Coccoderus to 15.

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DOI: 10.3897/asp.84.e176700

Authors: Jonas Eberle, Raphael Schallegger, Derek S. Sikes, Michel Lebel, Roland Gerstmeier

Abstract: Abstract The North American checkered beetle (Cleridae) Thanasimus nubilus Klug, 1842 is currently listed as a subspecies of Thanasimus undatulus (Say, 1835). We test the species status of T. nubilus by investigation of DNA barcodes and morphological measurements of multiple specimens of each of the two taxa as well as of multiple specimens of two additional North American species of the genus. We use four species delimitation methods based on gene tree and clustering algorithms (barcode gap threshold clustering, Assemble Species by Automatic Partitioning [ASAP], and single and multi-rate Poisson Tree Processes [mPTP]). Evidence from mtDNA clearly supported treating these two subspecies as distinct species which was also supported by a morphometric analysis. We therefore propose Thanasimus nubilus Klug, 1842 stat. rev. to be reinstated as a valid species.

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DOI: 10.3897/asp.84.e162254

Authors: Alberto Sánchez-Vialas, Arnau Calatayud-Mascarell, José L. Ruiz, Ernesto Recuero, Mario García-París

Abstract: Abstract The evolutionary dynamics of morphological traits can often blur the boundaries between interspecific divergence and intraspecific variability, complicating species recognition. This study investigates the variation in secondary sexual traits and the existence of potential speciation processes within what is now considered Berberomeloe insignis (Coleoptera: Meloidae), an endangered blister beetle taxon endemic to southeastern Spain. Despite previous evidence of substantial genetic and phenotypic differentiation, key characters as the morphological variation in secondary sexual traits, such as antennomeres, remain unexplored. Using geometric morphometrics, we analyzed the shape variation of male and female antennomeres VII–XI across all previously recognized lineages of B. insignis. Our results reveal significant morphological differentiation, particularly in antennomeres VII, IX, and XI, which correlate broadly with genetic lineages. Based on the study of newly recorded populations, we confirm that cephalic coloration patterns correspond with mitochondrial lineages, further supporting the existence of geographic lineage differentiation within what was previously considered B. insignis. Climatic niche modeling indicates low climatic niche overlap between the isolated western lineage and the remaining lineages, which also show relatively low to moderate overlap, suggesting that ecological factors could have contributed to the divergence among them. These findings underscore the intricate interplay of genetics and ecology, highlighting the importance of integrating multiple data sources for accurate species delimitation. Based on our results, we describe B. nazari sp. nov. and B. insignis trisanguinatus ssp. nov. reflecting the evolutionary history of this group.

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DOI: 10.3897/asp.84.e179328

Authors: José Ricardo Assmann Lemes, Ricardo Russo Siewert, Olaf Hermann Hendrik Mielke, Mirna Martins Casagrande, Andrew David Warren

Abstract: The taxonomy of the subgenus Staphylus (Capilla) is reviewed, including redescriptions of known species, identification keys, and detailed distribution maps. The female genitalia of Staphylus (Capilla) azteca (Scudder, 1872), S. (C.) caribbea (Williams & Bell, 1940), S. (C.) corumba (Williams & Bell, 1940), S. (C.) eryx Evans, 1953, S. (C.) tucumanus (Plötz, 1884) and S. (C.) tyro (Mabille, 1878) are described and illustrated for the first time. Four new species are described: Staphylus (Capilla) nicoleae Lemes sp. nov. from Colombia and Venezuela, S. (C.) ricardoi Lemes, sp. nov. from Peru, S. (C.) neideae Lemes sp. nov. and S. (C.) neivae sp. nov. from Brazil. Pholisora imperspicua Hayward, 1940 is a syn. nov. of Staphylus (C.) lizeri lizeri (Hayward, 1938) and Hesperia melangon epicaste Mabille, 1903 is a syn. nov. of Staphylus (C.) melangon melangon (Mabille, 1883). Neotypes are designated for Staphylus epicaste Mabille, 1903, Nisoniades tucumanus Plötz, 1884 and Staphylus fascia Hayward, 1933. Lectotypes are designated for Pholisora azteca Scudder, 1872, Bolla machuca Schaus, 1913, Helias tyro Mabille, 1878, Staphylus anginus Schaus, 1902, Hesperia melangon 1883, Hesperia musculus Burmeister, 1875 and Helias aurocapilla Staudinger, 1876.

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DOI: 10.3897/asp.84.e174748

Authors: Guanliang Meng, Leonardo S. Carvalho, Lars Podsiadlowski, Bernhard A. Huber

Abstract: Pholcid spiders have long been classified into five subfamilies, and this framework ultimately dates back to Eugène Simon’s non-phylogenetic system of 1893. While subfamily relationships and compositions have been updated extensively over the last decades, no new subfamily had to be erected for any of the hundreds of new species newly described since Simon. Here we report two new species from semi-arid Brazilian Caatinga: Caipira mineira Huber sp. nov., and Caipira baiana Huber sp. nov. Genomic data strongly support their sister-group relationship; we thus join them conservatively in a single genus, Caipira Huber gen. nov., even though they show some remarkable morphological differences. This genus is sister to a large clade including all pholcid subfamilies except Pholcinae and Smeringopinae, which necessitates the erection of a new subfamily: Caipirinae subfam. nov. In addition, we formalize the separation of the genus Artema from ‘other Arteminae’. This had previously been suggested by multi-locus genetic data, and is strongly supported by new genomic data. Arteminae is newly circumscribed to include only Artema and Priscula, and the name Physocyclinae subfam. nov. is proposed for ‘other Arteminae’. Pholcidae is thus divided into seven subfamilies, with the following relationships suggested by genomic data: (Pholcinae, Smeringopinae), (Caipirinae, ((Arteminae, Ninetinae), (Physocyclinae, Modisiminae))). Finally, we tested the hypothesis that the Chilean genus Aucana is the closest relative of the new Brazilian species. This is strongly rejected; Aucana is resolved as the only known South American representative of Physocyclinae.

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DOI: 10.3897/asp.84.e161376

Authors: Anna A. Namyatova, Polina A. Dzhelali, Darya S. Bolshakova

Abstract: Apolygus (Insecta: Heteroptera: Miridae: Mirinae) is a large and taxonomically challenging trans-Palearctic genus comprising 74 species, and it has never been revised. The greatest diversity of this genus occurs in Asia, and 15 species have been recorded from Russia. Among them, Apolygus limbatus, Apolygus lucorum, and Apolygus spinolae are trans-Palearctic, and A. lucorum and A. spinolae are economically important as pests. Previous studies have shown that A. lucorum and A. spinolae are very similar in morphology and that the barcoding region might not be suitable for their identification. In this study a revision of the Apolygus representatives inhabiting Russia is provided based on morphological and molecular data. Mitochondrial (COI and 16S rRNA) and nuclear markers (ITS1 and 28S3 rRNA) were used for phylogenetic analyses and automatic species delimitation methods were applied. Apolygus lucorum, A. malaisei, A. nigronasutus, A. nigrovirens were treated as junior synonyms of A. spinolae because morphological and molecular data did not show differences between those species. Apolygus shikotan sp. nov. was described as new to science. Morphological data and the COI phylogeny show that A. limbatus might represent two separate species. However, additional specimens and nuclear markers for both groups are needed to test this hypothesis. The synonymy of A. syringae with A. hilaris was supported based on the morphological data. Apolygus gotorum and A. subhilaris are reported from Russia for the first time. The nuclear marker ITS1 is the most reliable marker for species delimitation and identification of Apolygus species.

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DOI: 10.3897/asp.84.e162008

Authors: Sumeyra Nur Sanal Demirci, Volkan Kılıç, Serap Mutun, A. Yavuz Kılıç

Abstract: Tabanus bifarius Loew, 1858 is a horsefly species distributed throughout the Mediterranean region and in the areas of central and southeastern Europe and partly in Asia. In this study, we conducted the first comprehensive phylogeographic and population genetic analysis of T. bifarius collected from 13 localities across Türkiye and Iran, through utilizing mitochondrial (COI) and nuclear DNA (ITS1–5.8S–ITS2) markers. A total of 81 haplotypes and 59 nuclear alleles were identified among 187 sequenced individuals. Both mitochondrial and nuclear DNA patterns and diversity observed in T. bifarius are indicative of complex historical and demographic processes. Among all the populations studied, Eskişehir and Hatay exhibited the highest genetic diversity, which may be due to the region’s topography and transitional zones. Phylogenetic analyses suggested that T. bifarius split from the outgroup species approximately 5.53 million years ago (MYA) during the Late Miocene–Early Pliocene, likely driven by tectonic and climatic events. Subsequent diversification events that occurred during times of climatic and environmental fluctuations in the Late Pliocene and Early Pleistocene also seemed to have significantly affected the species and gave rise to the formation of some important genetic lineages. Our analyses results indicate that T. bifarius exhibits a structured genetic landscape shaped by multiple refugial routes, geographic barriers, and Quaternary climatic oscillations. Further, our findings suggest that the species likely entered Anatolia through three distinct routes: (1) from the Levant region into southern Anatolia via Hatay; (2) from the Caucasus into northeastern Anatolia through Ardahan; and (3) from the Iranian Plateau into eastern Anatolia via Van.

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DOI: 10.3897/asp.84.e157646

Authors: Lucas Quevedo, Renato S. Capellari, Carlos José E. Lamas

Abstract: Understanding the biogeographic patterns that explain species distribution in the Americas is a long and challenging endeavor. Insects represent a potentially promising group for these kinds of investigations, but deficits in basic taxonomic and systematic knowledge hinder the development of many studies. In this context, a cladistic analysis of the Pan-American genus Achradocera Becker (Diptera: Dolichopodidae: Diaphorinae) was conducted using a morphology-based matrix comprising 54 characters and 18 terminal taxa, including all 12 known species of the genus and six outgroup taxa. The analysis yielded two topologies under equal weighting and one under implied weighting of characters. Achradocera was recovered as monophyletic, and two species groups are proposed: the barbata group (Nearctic: A. barbata and A. arcuata) and the femoralis group (Neotropical: A. excavata, A. femoralis, A. meridionalis, and A. tuberculata). Biogeographic analyses were performed using the Geographically Explicit Event Model (GEM) method, which identified vicariance as the primary driver of speciation events in Achradocera, with founder events playing a secondary role. The results provide insights into the patterns that may explain the evolutionary history of the genus across the Americas, and potentially contributing to the understanding of patterns observed in other taxa with similar distributions, while also fostering discussions that can be integrated into broader frameworks, such as insect biogeography and studies of other taxa.

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DOI: 10.3897/asp.84.e153168

Authors: Xu-Hong-Yi Zheng, Yong Qin, Zhi-Teng Chen

Abstract: The genus Hexarthrius Hope, 1842, a prominent member of the family Lucanidae, is widely distributed across southern Asia and includes some of the largest stag beetle species in the world. This study presents the first mitogenomic phylogeny for Hexarthrius and its related genera, using complete mitogenomes from all recognized Hexarthrius species and representatives of all known related genera; 23 mitogenomes are reported here for the first time. We identified two novel mitochondrial gene rearrangements in Hexarthrius and its relative genus Rhaetulus Westwood, 1871, with implications for mitogenome evolution in Lucanidae. Phylogenetic inference and molecular dating recover Hexarthrius as monophyletic and subdivided into two well-supported clades: a Himalayan clade and a Tropical clade, which diverged approximately 8.9 million years ago. Rhaetus Parry, 1864 is inferred as the sister group to Hexarthrius, whereas Rhaetulus occupies a basal position within the larger Hexarthrius clade. Ancestral-area and morphological reconstructions indicate a complex history of vicariance and dispersal associated with uplift of the Hengduan–Himalayan region and subsequent island isolations. These results clarify species relationships, biogeography, and morphological evolution within this emblematic beetle lineage.

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DOI: 10.3897/asp.83.e158227

Authors: Ana Lucia Henriques-Oliveira, Jorge Luiz Nessimian, Daniela Maeda Takiya, Allan Paulo Moreira Santos

Abstract: Triplectides Kolenati, 1859 (Leptoceridae) is the most diverse genus within Triplectidinae, with about 90 species worldwide, 18 of them in the Neotropics. Currently, eleven species are recorded from Brazil. Since Holzenthal’s 1988 revision, many additional specimens have become available, leading to a significantly broader known distribution for several Brazilian species of Triplectides. Consequently, the morphological boundaries between some species have become less distinct. Furthermore, despite the ecological importance of caddisfly larvae, the immature stages of only a few Brazilian species have been described to date. The present study has the following aims: to evaluate Triplectides species boundaries using morphology and DNA sequences (COI and EF-1α); describe probable new species; and use DNA-based life stages associations to recognize and describe larvae at the species level. We delimited and formally described seven new species of Triplectides: T. bandeira sp. nov., T. caparaoensis sp. nov., T. cerradoensis sp. nov., T. iguassu sp. nov., T. mantiqueira sp. nov., T. paragracilis sp. nov., and T. puri sp. nov. Additionally, we successfully associated adult males with the immature larval stages of seven Brazilian species, including both newly described and previously known species of Triplectides, for which we provide morphological descriptions.

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DOI: 10.3897/asp.83.e159694

Authors: Shodo Mtow, Ryuichiro Machida

Abstract: We examined the morphology of the male and female genitalic regions of Petrobiellus akkesiensis and describe its mating behavior as analyzed using video recordings. Petrobiellus belongs to the monogeneric subfamily Petrobiellinae (Machilidae), an enigmatic group known only from the Far East. Its male genitalic region shows remarkable specialization, leading to the inference that Petrobiellus performs direct spermatophore transfer through genital coupling. Video analysis revealed that, as expected, the male holds the female’s ovipositor using both his penis and styli on the 9th abdominal segment: the penis grasps the basal region of the ovipositor with its hook-like tip and the styli hold the midsection of the ovipositor from above and below; the spermatophore is then discharged onto the dorsal side of the basal region of the ovipositor from the apex of the penis, and the sperm is taken into the ovipositor, thereby completing the direct sperm transfer from male to female. This is the first documented case of direct sperm (spermatophore) transfer via genital coupling in apterygote hexapods. Based on current and previous findings, we compared and characterized the mating behaviors of Archaeognatha and discuss their implications in terms of evolution of mating strategies in Hexapoda. The mating behavior of Petrobiellus represents one of the most derived modes within Machilidae and Archaeognatha, for which we propose the name “direct transfer of spermatophore by genital coupling”.

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DOI: 10.3897/asp.83.e171454

Authors: Josemária Silva de França, Bruno Cavalcante Bellini, Nerivânia Nunes Godeiro, Nikolas Gioia Cipola

Abstract: Rhynchocyrtus Mendonça and Fernandes, 2007 is a monotypic genus of Entomobryidae, endemic to Brazil. Its placement within the Lepidocyrtinae and its systematic affinities with other members of the subfamily, especially with the subgenera of Lepidocyrtus Bourlet, 1839, have never been tested before. Here, we described the morphology and mitogenome of a new species of Rhynchocyrtus from the northeastern Brazilian Atlantic Forest (Rhynchocyrtus cleideae sp. nov. holotype female deposited in CC/UFRN: Brazil, Rio Grande do Norte State, Natal municipality), depicting for the first time the dorsal trunk chaetal pattern, homology and body pseudopores distribution for the genus. The new species description provided further data which we used to update the genus diagnosis, following the current standards used for other Entomobryidae. We also evaluated the phylogenetic placement of the genus within Lepidocyrtinae, testing its affinities with different subgenera of Lepidocyrtus. Our results point to Rhynchocyrtus as an ingroup of Neotropical Setogaster Salmon, 1951 subgenus, and not related to Cinctocyrtus Yoshii and Suhardjono, 1989 as previously suspected. Setogaster is likely a paraphyletic taxon, suggesting that some features currently used to separate Lepidocyrtus subgenera do not hold phylogenetic signal, and should be reevaluated. We discuss the problematic systematics of Lepidocyrtinae and reinforce the usefulness of some alternative morphological traits to better define its subgroups, based on the current knowledge of the group.

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DOI: 10.3897/asp.83.e171040

Authors: Maite Allegue, Nicolás Peralta-Seen, Nelson Ferretti

Abstract: The present study contributes to the understanding of the diversity within the tarantula genus Euathlus Ausserer, 1875. Through a combination of cladistic, molecular, and morphological analyses, three new species are described: Euathlus basalticus sp. nov. from Neuquén, Euathlus kupal sp. nov. from Mendoza, and Euathlus susanae sp. nov. from the La Pampa-Mendoza provinces. Additionally, the female of Euathlus tenebrarum Ferretti, 2015 is described for the first time, revealing that the previously attributed female of this species belongs to E. basalticus sp. nov. Phylogenetic trees from both morphological and molecular datasets are presented. Preliminary molecular analyses reveal the identity and support the proposal of the species treated here. Morphological analyses found considerable diversity within Euathlus, particularly in genitalic structures, challenging the general assumption of morphological homogeneity among mygalomorph spiders. Cladistic analyses recovered the genus Phrixotrichus within Euathlus, though this hypothesis could not be tested with molecular data due to the lack of available sequences for Phrixotrichus. These results suggest that future taxonomic revisions may be complemented once molecular data become available. Altogether, our results provide new insights into the systematics and diversity of Euathlus and highlight the importance of integrative approaches for unraveling evolutionary relationships within this group.

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DOI: 10.3897/asp.83.e147402

Authors: Qing-Bo Huo, Xiao Yang, Abdur Rehman, Yu-Zhou Du

Abstract: The phylogenetic relationship of Perlodidae is presented in this paper based on mitochondrial genomes with different matrix (PCG, PCGR, PCG12, PCG12R). This study includes molecular data from seven genera and seven species, including three rare genera and five species from China. The results show that the monophyly of subfamilies and tribes of Perlodidae is well supported, as well as that Perlodinae and Isoperlinae diverged from a common ancestor at the same time, while the split between Arcynopterygini and Perlodini + Diploperlini occurred after the split between the two subfamiles. A discussion of the origin and migration of the Holarctic and Palearctic-Oriental perlodid genera is provided in this paper.

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DOI: 10.3897/asp.83.e152233

Authors: Tácio Duarte, Pitágoras C. Bispo, Pablo Pessacq

Abstract: Abstract The Gripopterygidae family, a diverse group of stoneflies (Plecoptera) endemic to the Southern Hemisphere, has traditionally been divided into five subfamilies, though the monophyly of most remains uncertain due to limited morphological and molecular support. Here we conducted a morphology-based cladistic analysis using 50 characters and 41 taxa, including representatives from all five Gripopterygidae subfamilies and three Austroperlidae species, to test the monophyly of Gripopteryginae and determine the phylogenetic position of a newly discovered species, Tupiperla furcata sp. nov. The analysis, rooted with Penturoperla barbata (Austroperlidae), employed parsimony with implied weighting and tree bisection reconnection methods. Results supported a core Gripopteryginae clade, with absence of a posterior sclerite in tergum 10, but excluded Neopentura semifusca, which was more closely related to Antarctoperlinae. Additionally, Paragripopteryx munoai presented morphological divergence, suggesting it may require reclassification into a new genus. These results challenge the current subfamily classifications, particularly Dinotoperlinae and Leptoperlinae, and highlight the need for further taxonomic revision. To advance the understanding of Gripopterygidae phylogeny, we suggest incorporating molecular data and expanding taxon sampling throughout the Southern Hemisphere. Such efforts would clarify evolutionary relationships and biogeographic patterns, paving the way for a more robust classification of Plecoptera. Un resumen traducido al español puede consultarse en el suplemento electrónico (File S1).

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DOI: 10.3897/asp.83.e166986

Authors: Gideon Pisanty, Rémi Santerre, Teresa Martin, Sophie Cardinal, Thomas J. Wood

Abstract: Andrena (Micrandrena) Ashmead is the largest subgenus of Andrena Fabricius 1775, with more than 150 species distributed throughout the northern hemisphere. The subgenus has gained notoriety as one of the most difficult taxonomic groups of bees, and remains poorly studied across large parts of the Palaearctic region. Based on extensive collecting in Israel, Lebanon and Cyprus, together with efficient DNA barcoding and and re-examination of old museum material, we present a thorough revision of the species of A. (Micrandrena) occuring in the region of the Levant (Israel, the West Bank, Jordan, Lebanon and Syria) and the island of Cyprus, including a detailed identification key, and a summary of our knowledge of the distribution, phenology and foraging biology of each taxon. Our study enumerates forty-two species of Andrena (Micrandrena) from the Levant and Cyprus, including four species new to the region, and nine species new to science: Andrena aphroditae Pisanty sp. nov., A. alshaykh Pisanty sp. nov., A. aspera Pisanty & Wood sp. nov., A. chananaea Pisanty & Wood sp. nov., A. friedmani Pisanty sp. nov., A. hebraica Pisanty & Wood sp. nov., A. kugleri Pisanty sp. nov., A. libanica Wood sp. nov., and A. phoenicia Pisanty sp. nov. We describe the males of A. calandra Warncke, 1975 and A. lindbergella Pittioni, 1950 for the first time. We additionally synonymise Andrena dargia Warncke, 1965 with A. minutula (Kirby, 1802) syn. nov., and recognise two previously subspecific names as valid species: Andrena leptura Warncke, 1974 stat. nov. and A. povolnyi, 1974 Warncke stat. nov.

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DOI: 10.3897/asp.83.e145082

Authors: Yuchen Zheng, Yuezheng Tu, Davide Badano, Xingyue Liu

Abstract: Abstract Layahima Navás, 1912 is the most diverse antlion genus of the tribe Acanthoplectrini (Myrmeleontidae: Dendroleontinae) endemic to the Oriental region, currently comprising 12 species. However, the species diversity of this genus is still far from completely explored, and its larval stage is poorly known. Here, we describe four new species of Layahima, i.e., L. haohani sp. nov., L. qilin sp. nov., L. pixiu sp. nov., and L. zhitengi sp. nov., from Southwest China. Moreover, we describe the larval stages of three Layahima species, i.e., L. chiangi Banks, 1941, L. lhoba Zheng, Badano, Liu, 2023, and L. yangi Wan & Wang, 2006. The precise distribution of L. chiangi, whose type locality was previously unclear, has now been clarified to be exclusively restricted to the Nujiang dry hot river valley around Cawarong, Xizang. The phylogeny of Layahima by adding new species herein reported was inferred based on molecular data. The L. zonata group, once considered monophyletic, was recovered as paraphyletic within Layahima.

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DOI: 10.3897/asp.83.e164315

Authors: Felipe Francisco Barbosa, André Silva Roza, José Ricardo M. Mermudes, Michael F. Geiser, Jiri Hodecek, Lara-Sophie Dey, Michael A. Ivie, Viridiana Vega-Badillo, Vinicius S. Ferreira, Robin Kundrata

Abstract: Abstract Phengodidae, or railroad-worm beetles, are bioluminescent soft-bodied beetles with flight capable adult males and highly paedomorphic larviform females. They are well accepted as part of the “lampyroid” clade within Elateroidea, and their suprageneric relationships have been recently studied using a phylogenomic approach. However, the placement of taxa currently classified in the subfamily Penicillophorinae remained untested. Penicillophorinae form an assemblage of morphologically modified, rarely collected Neotropical genera that are unique among Phengodidae. They are particularly characterized by their moniliform, serrate or uniramose antennae, in contrast to the typically bipectinate antennae present in all other members of the family. To investigate the phylogenetic position of Penicillophorinae, we implemented a low-coverage whole genome sequencing approach to produce genomic data for Acladocera and Walterius, two out of five genera classified in this subfamily. The resulting phylogenomic analyses confirmed the monophyly of Phengodidae; however, Penicillophorinae were not found as a monophyletic group. Our results recovered the topology: Cydistinae + (Phengodinae + (Acladocera + (Cenophenginae + Mastinocerinae including Walterius))). Therefore, we suggest that the antennal double rami were lost at least twice among Phengodidae: once in the newly circumscribed Mastinocerinae (with Walterius), and once in Acladocera, which we tentatively keep in Penicillophorinae. Further, we discuss the morphological modifications of other genera currently classified in Penicillophorinae. Future phylogenomic research should focus on clarifying the boundaries and composition of phengodid subfamilies, particularly by including additional genera from Penicillophorinae and Mastinocerinae.

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DOI: 10.3897/asp.83.e161465

Authors: Mathieu Boderau, Matteo Montagna, Fabio Magnani, Giulia Magoga, Alexey Shmakov, Michael S. Engel, Patricia Nel, André Nel

Abstract: Abstract We report the discovery of the first Triassic ‘shielded thrips’ of the family Iotacyphidae, represented by two species: Triassocypha prima gen. et sp. nov. and T. secunda sp. nov. The species are described and illustrated, and comparisons are made with other stem-Thysanoptera lineages. The unique wing venation of Iotacyphidae and related families is reinterpreted in the light of these new fossils. The venation pattern supports the diagnosis of the family, as well as of higher clades within the broader lineage that includes both thrips and their extinct relatives (in the total group Holothysanoptera, formerly Thripida), and contributes to clarifying the systematics of thrips.

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DOI: 10.3897/asp.83.e144844

Authors: Kirstern Lica Follmann Haseyama, Claudio José Barros de Carvalho, Elaine Della Giustina Soares, Silvio Shigueo Nihei

Abstract: Abstract Polietina is a genus of Muscidae found in the New World, particularly in the Neotropical region. The genus is currently classified within the subfamily Muscinae and consists of 15 species whose phylogenetic relationships have been previously studied using morphological data. In our study, we conducted a phylogenetic analysis based on 37 morphological characters sourced from the literature and eight molecular markers (12S, 16S, 18S, 28S, cytochrome C oxidase subunit I, cytochrome b, region 4 of carbamoyl-phosphate synthetase-aspartate transcarbamoylase dihydroorotase, and elongation factor 1-ɑ), totaling 5366 characters. We obtained molecular data for eight species of the genus and four outgroup taxa from original sequencing and public repositories. We used Bayesian posterior probabilities to estimate the topology, as follows: (P. flavidicincta ((P. rubella (P. concina (P. wulpi, P. orbitalis))) (P. steini ((P. flavithorax (P. ponti, Polietina sp.)) (P. major (P. prima (P. bicolor (P. minor (P. univittata, Polietina sp.))))))))). The reciprocal monophyly of all species for which more than one identified sequence was available is supported by our results. Our analysis largely supports the previously published hypotheses regarding the phylogeny of Polietina. However, major differences were observed between the locations of P. flavidicincta and P. flavithorax. Additionally, we discuss the identity of unidentified Polietina specimens with sequences published in GenBank or new sequences produced as part of our study.

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DOI: 10.3897/asp.83.e151385

Authors: Bernardo F. Santos, Santiago Bordera

Abstract: Abstract The phylogeny of the Old World genus Stenarella Szépligeti is investigated using data from genomic ultraconserved elements. Thirteen species from the four biogeographic regions where the genus occurs were included in the analyses. The resulting trees corroborate the monophyly of Stenarella and show substantial biogeographic structure within the genus, with three major clades corresponding to species occurring in the Afrotropical, Palearctic and Oriental–Australasian regions. The Afrotropical component of the genus is represented by fifteen species, while outside of the Afrotropics there seem to be only three species, each with very wide geographic distribution. The taxonomic limits of the genus are reviewed, and an updated diagnosis is provided. A total of eighteen species are recognized as valid, of which twelve are described as new: S. aurea sp. nov., S. bugalana sp. nov., S. delicata sp. nov., S. favilla sp. nov., S. fenestralis sp. nov., S. hopkinsi sp. nov., S. katanga sp. nov., S. natalina sp. nov., S. nigriscuta sp. nov., S. nigromaculata sp. nov., S. saaksjarvi sp. nov., and S. vannoorti sp. nov. An updated identification key to the species of the genus is provided, as well as diagnoses, redescriptions, and new distribution records for the previously known species.

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DOI: 10.3897/asp.83.e146637

Authors: Danilo César Ament, Eduardo A.B. Almeida

Abstract: Abstract We address an old but still controversial question of morphological phylogenetics: whether additive (or ordered) coding is beneficial to properly extracting phylogenetic information from phenotypical variation. To empirically evaluate the value of the additive coding, we compared the impact of multistate additive, non-additive, and binary codings for 14 quantitative characters in a phylogenetic analysis of a genus of phorid flies (Diptera). First, we compared which of these morphological codings were most effective for the morphological matrix to approximate the results of a molecular data set. We then compared which morphological coding strategies yielded the best Bayesian posterior probabilities when concatenated to molecular data. We also calculated consistency and retention indices for each binary element of the additive characters and contrasted these results to a measure of phylogenetic signal. Overall, these indices were lower for additive characters than for the others but still indicate reasonable accommodation in the tree. Additive coding outperformed the multistate non-additive coding by recovering higher Bayesian posterior probabilities in the concatenated dataset. Additive coding was also among the best coding strategies for the morphological matrix to approximate the phylogenetic signal from an independent source of evidence—i.e., molecular results. Therefore, quantitative information coded as additive had reasonable phylogenetic congruence with other data and improved the phylogenetic results of morphological data in most cases. These results support the use of additive coding for phylogenetic analysis and encourage other similar empirical evaluations aiming to explore the generality of the benefits of this coding method.

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DOI: 10.3897/asp.83.e150460

Authors: Daniele Camarda, Oscar Lisi, Daniel Stec, Matteo Vecchi

Abstract: Abstract A new tardigrade taxon, Thulyphoribius melitense gen. nov. et sp. nov. is described from a population of limnic tardigrades collected in the sediment of a temporary pond in Malta. Those habitats, characterized by fluctuating environmental conditions and ephemeral water availability, provide a particularly challenging setting that can drive morphological and genetic diversity in aquatic microfauna. The new genus shows a unique combination of morphological characters, including distinctive morphologies of the peribuccal region, a short and wide ventral lamina and Pseudobiotus-type claws. Although it shares certain traits with some extant genera, the presence of unique characters combinations, precludes its assignment to any of the previously established genera. A comprehensive investigation was conducted, including morphological (using Phase-Contrast Light Microscopy, Scanning Electron Microscopy), morphometric, and molecular analysis. In the latter context, the molecular markers 18S, 28S, COI, and ITS2 were sequenced, and the complete mitochondrial genome was obtained and characterized, offering important insights for future molecular studies of tardigrades. Phylogenetic analyses based on Maximum Likelihood and Bayesian methods, incorporating the four aforementioned molecular markers, indicate that the new genus belongs to the superfamily Isohypsibioidea, yet it does not cluster within any currently recognized extant family of tardigrades. Collectively, these findings underscore the relevance of investigating underexplored habitats and highlight the potential for discovering novel evolutionary lineages among limnic microinvertebrates that inhabit temporary ponds.

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DOI: 10.3897/asp.83.e153226

Authors: Emma Kärrnäs, Christer Hansson, Niklas Wahlberg

Abstract: Abstract Despite their ecological and economic importance as key regulators of insect populations worldwide, eulophid wasps remain one of the taxonomically most neglected groups of insects. As these minute parasitoid wasps exhibit an astonishing species diversity while being challenging to identify, relying on morphological characters alone has long been problematic. By using large-scale barcoding and molecular species delimitation analyses of 909 specimens, of which 641 were of European origin, we reveal a remarkable diversity of cryptic species in the two eulophid wasp genera Oomyzus Rondani and Quadrastichus Girault (Eulophidae, Tetrastichinae) present in Europe. We also present 40 new country records for 17 species and greatly improve the existing DNA barcode libraries of these two groups. Using the four different species delimitation methods ABGD, ASAP, GMYC and mPTP, the number of potentially undescribed European species detected in this study ranges from 20 to 34, of which 16 to 24 are cryptic. These results suggest that not even half of the European Oomyzus and Quadrastichus species are currently known, thus highlighting the need for further taxonomic work. However, the results also highlight issues with molecular species delimitations based on barcode data, such as incongruence across different analysis methods and potential barcode sharing between closely related species. Therefore, the results also challenge the validity of relying on a single molecular marker when detecting and subsequently describing new species, an increasing practise within modern taxonomy termed “turbo-taxonomy”.

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DOI: 10.3897/asp.83.e155906

Authors: Sixun Ge, Tai Gao, Yizhou Liu, Jiale Li, Juan Shi, Lili Ren, Hongliang Shi

Abstract: Abstract Sirex wood wasps (Hymenoptera: Siricidae) constitute a group of economically significant xylophagous pests that cause substantial damage to forestry globally. Herein, we present a systematic revision of the Chinese Sirex species. Through phylogenetic analyses, some taxonomic ambiguities were resolved. Sirex chimera sp. nov. is described (type locality: Chifeng, Inner Mongolia). Two new synonyms are proposed: S. rufiabdominis Xiao & Wu, 1983 and S. sinicus Maa, 1949 are synonymized with S. nitobei. S. dux (Semenov, 1921) stat. rev. is confirmed as a distinct species rather than a subspecies of S. juvencus or S. cyaneus, with the male described for the first time. Re-examination confirms that previous Chinese records of S. imperialis represent misidentifications of S. nitobei. The Chinese Sirex fauna is revised to comprise nine valid species: S. chimera sp. nov., S. dux, S. ermak, S. juvencus, S. nitobei, S. noctilio, S. piceus, S. tianshanicus, and S. vates. Comprehensive re-descriptions and a key to species are provided.

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DOI: 10.3897/asp.83.e151555

Authors: Fang-Shuo Hu, Alexey Solodovnikov, Martin Fikáček

Abstract: Abstract We provide a taxonomic description of the larva of the rove beetle Tolmerinus fratrumelliotorum Rougemont, 2017, identified through DNA barcoding. This is the first larval description for Anisolinina, the only subtribe whose sister group within the tribe Staphylinini remains unclear. While most phylogenetic hypotheses, especially molecular-based, suggest sister group relationships between Anisolinina and Staphylinina, conflicts remain, particularly when adult morphology is used for phylogeny reconstruction. Our attempt to use larval morphology for phylogeny reconstruction neither revealed a sister group for Anisolinina, nor unique synapomorphies for this subtribe. However, by mapping larval characters on the reference tree of Staphylinini, i.e., a phylogeny firmly established by phylogenomics in agreement with the adult morphology, we found unique larval synapomorphies to support the tribe Staphylinini and informal clade Staphylinini propria. We review all phylogenetic hypotheses ever proposed for a sister group of Anisolinina and discuss potential reasons for the poor phylogenetic signal in the larval characters alone. We stress some opportunities and challenges of using larval characters in phylogeny reconstructions.

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DOI: 10.3897/asp.83.e152943

Authors: Álvaro Laborda, Miguel Simó, Luis N. Piacentini, Antonio D. Brescovit, Carolina Beloso, Anita Aisenberg, Miquel A. Arnedo, Martín J. Ramírez, Leticia Bidegaray-Batista

Abstract: Abstract The wolf spiders of the subfamily Allocosinae are known for their complex taxonomy, especially in the Neotropical region. Despite previous taxonomic and phylogenetic studies, the diversity and phylogenetic relationships of the subfamily remain largely unknown. This study aims to clarify the evolutionary relationships within South American Allocosinae, hypothesizing a greater diversity than currently recognized and seeking to resolve ambiguities in genus-level classification. We used a combination of mitochondrial and nuclear gene sequences to construct phylogenetic analyses for 73 specimens across 13 species of Allocosinae. Analyses using both maximum likelihood and Bayesian frameworks were conducted to examine internal relationships and phylogenetic structure and to infer a timeline of diversification. Additionally, species delimitation was conducted to identify cryptic lineages. Our results recover the specimens considered to be representatives of the subfamily Allocosinae as a monophyletic group, and identified five major clades. Divergence time estimates suggested Allocosinae originated in the Early Miocene (15–22 million years ago), and underwent significant diversification during the Pleistocene. Species delimitation analysis based on single markers uncovered 24 lineages, indicating potentially overlooked species. Allocosinae has shown to be an interesting group to study incipient speciation processes, ecology of coastal environments and atypical behaviors such as sex role reversal. Knowing and understanding the evolutionary history and relationships within the subfamily is necessary for progress in its study in any field of biology.

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DOI: 10.3897/asp.83.e150304

Authors: Mariana Raquel Chani-Posse, Maryzender Erceliz Rodríguez-Melgarejo, José Manuel Ramírez-Salamanca

Abstract: Abstract Konradus Chani-Posse & Ramírez-Salamanca, 2020 was originally described as a monotypic genus from the tropical Andes of Ecuador. However, a re-examination of type material from species previously classified under the genus Philonthus Stephens, 1829, along with additional, previously unstudied specimens collected from high-altitude regions (above 2000 m) in the Andes of Ecuador, Peru, and Bolivia, prompted a more comprehensive reassessment of the genus. This study presents a revision of Konradus and introduces Yuracarus gen. nov., based on external morphology and male and female sexual characters. Two species previously considered “false Philonthus” (Ph. actinus Bernhauer, 1917 and Ph. diamantinus Bernhauer, 1917) are transferred to Konradus and Yuracarus gen. nov., respectively: K. actinus (Bernhauer), new comb., and Y. diamantinus (Bernhauer), new comb. Additionally, two species from Peru are newly described within Konradus: K. cuscensis sp. nov. and K. trescrucensis sp. nov., while three new species are described within Yuracarus: Y. cosnipatensis sp. nov., Y. napoensis sp. nov., and Y. yunguensis sp. nov. A key to species, diagnoses, descriptions and/or redescriptions, illustrations, a distributional map, and a phylogenetic analysis are included. Lectotypes are designated for Philonthus actinus Bernhauer and Philonthus diamantinus Bernhauer. Cladistic analysis confirms the monophyly of Konradus and Yuracarus within the Andean clade (AC) of Neotropical Philonthina and supports Yuracarus as a distinct genus. Both genera are part of a well-supported clade characterized by the presence of sub-bilobed and subtriangular protarsomeres 2 and 3, bearing discal setae on their ventral surface, as well as a horseshoe-shaped accessory sclerite associated with female tergum 10.

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DOI: 10.3897/asp.83.e153079

Authors: Edvanda A. Souza-Carvalho, Célio Magalhães, Fabrício L. Carvalho, Fernando L. Mantelatto

Abstract: Abstract The generic system of the Neotropical crabs of the subfamily Trichodactylinae H. Milne Edwards, 1853 has remained quite stable over the last 30 years, but the recognition or not of the genus Mikrotrichodactylus Pretzmann, 1968 has been a matter of debate: erected as a subgenus, it was treated with generic status in Rodríguez’ classificatory system but some subsequent works considered it a junior synonym of Trichodactylus Latreille, 1828. Based on this scenario, an integrative analysis based on molecular (using two mitochondrial, 16S rRNA and COI, and one nuclear, Histone 3, genes) and morphological (using diagnostic characters traditionally used on the identification of the family) data was performed in order to clarify the phylogenetic position of the genera within Trichodactylinae. The inferred phylogeny recovered three great lineages within Trichodactylinae with high support values in both Bayesian Inference and Maximum Likelihood phylogenetic analyses, corroborated the non-monophyletic status of Trichodactylus, and confirmed the full generic status of Mikrotrichodactylus. A taxonomic rearrangement of Trichodactylinae is proposed but the positioning of “Trichodactylus” quinquedentatus Rathbun, 1893 remains doubtful and was treated herein as a taxon inquirendum et incertae sedis. This taxon seems to be closer related to Rodriguezia Bott, 1969, and Avotrichodactylus Pretzmann, 1968; however, more data are needed before additional taxonomic adjustments concerning its positioning within the subfamily is proposed. In the current proposal, Trichodactylus is composed by four species, Mikrotrichodactylus by six species, Avotrichodactylus and Rodriguezia by three each, in addition to “Trichodactylus” quinquedentatus.

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DOI: 10.3897/asp.83.e150736

Authors: Mariano C. Michat, Yves Alarie, Cesar J. Benetti, Juan I. Urcola, Georgina Rodriguez, Patricia L. M. Torres

Abstract: Abstract A comprehensive phylogenetic analysis of larval characters, including for the first time members of the rare and enigmatic genus Queda Sharp, 1882 was conducted to test the hypotheses of monophyly and relationships of the tribe Hydrovatini within the diving-beetle subfamily Hydroporinae. Our results indicate that Hydrovatini (including both Queda and Hydrovatus Motschulsky, 1853) is monophyletic and unambiguously supported by the absence of the primary pore PAo on the parietal and by the antennal process (commonly identified as A3’) arising from the base of the antennomere 4. This result agrees with the original concept of the tribe introduced more than 100 years ago but challenged by some authors. Our study supports the inclusion of Hydrovatini in a clade of ancestral hydroporine lineages together with Laccornini, Laccornellini and Pachydrini. Within this clade, Hydrovatini is sister to Pachydrini based on the shared absence of an occipital suture in instars II–III, although with weak support. The third-instar larva of Queda is diagnosed and described. It is characteristic in the broad and semicircular shape of the nasale, the maxillary cardo partially fused to the stipes, the presence of setiferous tubercles on the cephalic capsule and abdominal segment VIII, and the subcylindrical and relatively well-developed galea which, interestingly, is somewhat more developed than those exhibited by other hydroporines.

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DOI: 10.3897/asp.83.e137316

Authors: Magdalena Rost-Roszkowska, Łukasz Chajec, Dalibor Stojanović, Dragan Antić

Abstract: The middle endodermal region of the digestive system (midgut) of arthropods is responsible for processes related to digestion but is also considered an organ participating in homeostasis maintenance. Therefore, many experimental studies, for example, related to the effect of various stressors on the organism, are conducted on the epithelium of this intestine. However, it is important to know the basic structure and ultrastructure of the midgut tissues. In myriapods (e.g., millipedes), the midgut has the form of a simple tube lined with a single layer of epithelium, surrounded by hepatic cells and visceral muscles. Considering the fact that millipedes can inhabit various terrestrial environments, feed on a variety of foods, and are important links in food chains, they can be considered good models for ecotoxicological studies. Thus, we selected eight species belonging to the family Julidae (order Julida) to investigate whether any distinct ground pattern for this organ appears within one millipede systematic group and whether it is possible to translate it into the general pattern of the midgut epithelium in millipedes: Leptoiulus sarajevensis (Verhoeff, 1898), Leptoiulus trilineatus (C. L. Koch, 1847), Cylindroiulus boleti (C. L. Koch, 1847), Megaphyllum bosniense (Verhoeff, 1897), Pachyiulus cattarensis (Latzel, 1884) and Pachyiulus hungaricus (Karsch, 1881) as representatives of the epigean fauna as well as Leucogeorgia longipes Verhoeff, 1930 and Leucogeorgia gioi Antić and Reip, 2020 as true cave-dwelling species. The study was performed using light and transmission electron microscopy. The results revealed a general pattern of all cells forming the midgut epithelium in Julidae, as well as the hepatic cells surrounding the midgut. Some small differences were observed, which are probably related to the type of food consumed.

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DOI: 10.3897/asp.83.e138238

Authors: Mattia Ragazzini, Roy M. J. C. Kleukers, Luc Willemse, Baudewijn Ode, Lara-Sophie Dey, Oliver Hawlitschek

Abstract: The Mediterranean Basin, recognized as a global biodiversity hotspot, harbors a remarkable diversity of grasshoppers, katydids, and crickets, many of which are endemic and potentially contain cryptic lineages. In this study, we generated a comprehensive dataset comprising 1,441 barcodes from 270 identified species within the Ensifera and Caelifera suborders. These were combined with existing data to form a dataset of 2,606 barcodes representing 351 species. We employed Maximum Likelihood (ML) topology reconstruction and applied five species delimitation methods (BIN, ABGD, ASAP, GMYC, and PTP) to detect potential incongruences between Operational Taxonomic Units (OTUs) and existing taxonomic classifications. Our analysis revealed that OTUs delimited by these methods corresponded to 71.39% of the evaluated species, with a notably higher congruence in Ensifera (88.53%) compared to Caelifera (52.15%). Across the dataset, we identified 54 lineages comprising cryptic species, indicating significant unrecognized diversity within these groups. Additionally, 21 instances of species being merged into consensus OTUs were observed, suggesting either the need for taxonomic revision or highlighting the limitations of current genetic markers. Among the methods tested, ABGD, particularly with the Kimura two-parameter model, was the most consistent with traditional taxonomy, yielding the highest consensus rates. In contrast, the PTP method exhibited the lowest consensus, often leading to an oversplitting of lineages. These findings underscore the complexity of species delimitation in recently radiated taxa and emphasize the importance of using multiple methodologies to accurately capture biodiversity, especially in regions characterized by a high prevalence of cryptic species.

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DOI: 10.3897/asp.83.e143754

Authors: Luísa Alasmar, Alexandre Cruz Domahovski, Rodney Ramiro Cavichioli

Abstract: In a morphological-based analysis using implied weighting for 23 taxa and 50 characters, we provided a phylogenetic study for Ciminius. The genus was recovered as monophyletic with a high support. We also provided a taxonomic revision, with description of three new species, increasing the diversity of Ciminius from seven to ten species: the Neotropical C. albolineatus, C. callosa, C. platensis, C. yana, C. autumnalis sp. nov., C. dissidens sp. nov., C. sesamum sp. nov., and the Nearctic C. hartii, C. sidanus, and C. taosus. A new monotypic genus, Arcanus gen. nov., recovered as sister to Ciminius is described. Tylozygus Fieber, the new genus, and Ciminius were recovered forming a clade here nominated as Articulate Stem Clade, due to its unique connective morphology amongst all Cicadellini. The immature stages of Ciminius sp. are described and a key to males, an occurrence map, and notes of parasitism and submacroptery are provided. We discussed the Ciminius relationships according to our analysis, the genus distribution, and the taxonomic problems about C. callosa.

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DOI: 10.3897/asp.83.e130334

Authors: Douglas J. Hilton, D. Andy Young, Liz Milla, Mengjie Jin, Stephen Wilcox, Qike Wang, Verena C. Wimmer, Jinny Chang, Henning Kallies, Andie Hall, Marina Watowich, Carly A. Busch, Jordan Wilcox, Aileen Swarbrick, Marlene Walter, Don Sands, Davina Paterson, David C. Lees, Marco F. Duretto, Adnan Moussalli, Mike Halsey, Axel Kallies

Abstract: Heliozelidae are a group of small monotrysian moths with a near world-wide distribution. While the Heliozelidae fauna of the Palaearctic and Nearctic is relatively well known, few studies have examined Heliozelidae in other regions of the world. If known, described species are leaf miners as larvae; however, the early biology of species outside of the Northern Hemisphere is poorly understood. Here, we describe a group of heliozelid moths that are specialised pollinators of the iconic plant genus Boronia Sm. (Rutaceae) from the south of Western Australia. Females of this group are characterised by the presence of a pollen-collecting structure on the dorsal side of the abdomen that is unique among known Lepidoptera. We propose that these moths are involved in a brood pollination mutualism with their species-specific host plant, where females lay eggs into and pollinate Boronia flowers, and larvae consume developing seeds. Molecular phylogenetic analyses using seven mitochondrial protein coding genes recovered a monophyletic group of pollinator species that belong to a larger group of Rutaceae-associated Australian Heliozelidae. The remainder of this group lack this pollen-collecting structure, providing insights into the evolution of pollination relationships. We resurrect the genus name Prophylactis Meyrick, 1897 stat. rev. and describe 15 new species based on a combination of morphological and molecular characters and host plant information: Prophylactis albiflorallax Hilton, Young & Kallies sp. nov., P. binbin Hilton, Young & Kallies sp. nov., P. clavatallax Hilton, Young & Kallies sp. nov., P. crassifoliallax Hilton, Young & Kallies sp. nov., P. crenulatallax Hilton, Young & Kallies sp. nov., P. gracilipax Hilton, Young & Kallies sp. nov., P. heterophyllax Hilton, Young & Kallies sp. nov., P jasperae Hilton, Young, Milla & Kallies sp. nov., P. megastigmallax Hilton, Young, Halsey, Milla & Kallies sp. nov., P. molloyax Hilton, Young & Kallies sp. nov., P. octandrallax Hilton, Young, Milla & Kallies sp. nov., P. pulchellax Hilton, Young & Kallies sp. nov., P. purdieanallax Hilton, Young & Kallies sp. nov., P. strictallax Hilton, Young, Halsey & Kallies sp. nov., and P. tetrandrallax Hilton, Young, Milla & Kallies sp. nov.

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DOI: 10.3897/asp.83.e137073

Authors: Hoi-Sen Yong, Sze-Looi Song, Kah-Ooi Chua, Yvonne Jing Mei Liew, I. Wayan Suana, Phaik-Eem Lim, Kok-Gan Chan, Praphathip Eamsobhana

Abstract: The complete mitogenomes of fruit flies Zeugodacus (Javadacus) calumniatus, Z. (Javadacus) heinrichi and Z. (Sinodacus) hochii have similar gene order and contain 37 genes and a non-coding region. They share an identical start codon for the respective protein-coding genes (PCGs), an identical TAA stop codon for 11 PCGs, TAG for cob, and an incomplete T stop codon for nad5. The cloverleaf structure of most of the tRNAs is similar in the three Zeugodacus species. Phylogenetic analyses reveal Z. (Parasinodacus) cilifer to be external to two main clades: (A) monophyletic subgenus Zeugodacus; and (B) subgenera Javadacus and Sinodacus. The present results indicate that the taxonomic status of some taxa needs clarification. Z. calumniatus is genetically very similar to Z. tau and is not congruent with its current placement in the munda complex. Z. mukiae NC_067083 is genetically very similar to Z. scutellaris, but differs significantly from Z. mukiae MG683384 of the arisanicus (arisanica) complex. On the other hand, Z. proprediaphorus is genetically distinct from and not a synonym of Z. diaphorus. Z. caudatus sensu stricto from Indonesia forms a sister lineage with Z. diversus, instead of with the Malaysian and Chinese taxa of Z. caudatus sensu lato. A notable incongruence is the sister lineage of Z. (Sinodacus) hochii and Z. (Javadacus) heinrichi among other taxa of subgenus Javadacus. A more extensive taxon sampling, particularly the subgenus Sinodacus (and other subgenera), is needed to clarify/resolve their subgenus status.

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DOI: 10.3897/asp.83.e143543

Authors: Jesús Gómez-Zurita, Anabela Cardoso

Abstract: In this work, we use a combined analysis of morphological and mtDNA sequence data to recognize and revise a group of species allied to the New Caledonian endemic leaf beetle genus Cazeresia Jolivet, Verma & Mille, 2005 of the Eumolpinae, considered monotypic before this work. We characterize and describe 20 new species allied to C. montana Jolivet, Verma & Mille, 2005 based on the recognition of morphological diagnostic traits and DNA-based species delimitation: C. australis sp. nov., C. clipeata sp. nov., C. corrugata sp. nov., C. globosa sp. nov., C. gracilis sp. nov., C. holosericea sp. nov., C. imperiosa sp. nov., C. impressicornis sp. nov., C. laevigata sp. nov., C. laticollis sp. nov., C. maquis sp. nov., C. ovata sp. nov., C. parentalis sp. nov., C. petitpierrei sp. nov., C. robusta sp. nov., C. spadicea sp. nov., C. subgeminata sp. nov., C. tibialis sp. nov., C. tricolor sp. nov. and C. wanati sp. nov. For C. globosa and C. spadicea we additionally describe the subspecies C. globosa altitudinalis ssp. nov. and C. spadicea bruna ssp. nov. We also propose transferring to this genus the species Thasycles humboldtiana Heller, 1916, Colaspis kanalensis Perroud, 1864, Dematochroma thyiana Jolivet, Verma & Mille, 2008 and Dumbea striata Jolivet, Verma & Mille, 2007, as C. humboldtiana (Heller) comb. nov., C. kanalensis (Perroud) comb. nov., C. thyiana (Jolivet, Verma & Mille) comb. nov. and C. striata (Jolivet, Verma & Mille) comb. nov., respectively. At present, the genus Cazeresia includes 25 species, the vast majority distributed in the southern part of Grande Terre in areas characterized by ultramafic soils and we speculate that the adaptation to these environmental characteristics in lowland areas may be ancestral in this lineage. Two thirds of the species are only known from their type locality, thus treated as potential microendemics, and most other have reduced ranges generally spanning few tens of kilometres. Finally, the degree of species sympatry or parapatry exhibited by Cazeresia is noteworthy, which in the absence of marked morphological differences among species suggests the possibility of the interplay of ecological mechanisms to minimize competition and exclusion.

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DOI: 10.3897/asp.83.e136516

Authors: Onur Uluar, Dragan P. Chobanov, Battal Çıplak

Abstract: A taxonomical practice without integration of systematics remains incomplete or produces incorrect definitions. To address this, we linked the taxonomy of the Poecilimon zonatus group to its systematics by examining its phylogeny and phylogeography. We used both mitochondrial and nuclear genes representing all species except P. varicornis. The mitochondrial matrix was subjected to phylogenetic analyses and species delimitation tests while phylogenetic signals in the nuclear data were assessed via haplotype network. Species delimitation tests suggested 12–16 species and the genetically divergent lineages were examined for phenotypic disparity. The results led to the following conclusions. (i) after the nomenclatural changes made here, P. zonatus group consists of 13 species, two of which are polytypic; P. salmani Sevgili syn. nov. is synonymized with P. ciplaki; the subspecies P. ciplaki denizliensis Kaya and P. zonatus datca Sevgili, Sirin, Heller & Lemonnier-Darcemont are elevated to species level as P. denizliensis Kaya stat. nov. and P. datca Sevgili, Sirin, Heller & Lemonnier-Darcemont stat. nov., respectively; P. boncukdagensis sp. nov., P. parazonatus sp. nov. and P. anisozonatus sp. nov., are described as new species, and P. datca montana subsp. nov. and P. denizliensis kizildagi subsp. nov. as new subspecies, (ii) the common ancestor of species corresponds to Mid Pleistocene Transition, suggesting the major climatic turnovers as the main evolutionary driver, (iii) the isolated relict ancestors on highlands evolved under similar selection pressures leading to a conserved phenotype, consequently, to limited morphological divergence despite their genetic differences; (iv) although morphological diagnosability is poor, each species meets the criteria of several species concepts.

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DOI: 10.3897/asp.83.e137570

Authors: Max Kasparek, Mira Boustani

Abstract: The distribution of the anthidiine bee identified as Anthidium undulatum Dours, 1873 extends from the French Mediterranean coastal region across the Adriatic coast and the eastern Mediterranean to the Caucasus and Iran. Examination of the DNA sequence of the mitochondrial COI gene (“barcoding unit”) of 25 specimens from throughout the distributional range revealed the existence of three distinct clades, which are clearly separated with bootstrap values of 99% and 100% in a Maximum Likelihood Analysis. The genetic distance between these groups ranged from 2.6 to 5.0%, while the intra-group distance did not exceed 0.42%. Two of these clades were found to occur in the same habitat in Lebanon. These three clades also differ in some phenotypic color traits, which, however, are not always fully diagnostic. Despite this, a morphometric analysis of six parameters of the head and wings of the males showed that the three genetic clades also form distinct clusters in a Discriminant Function Analysis. The consistent results of the genetic and morphometric analyses, together with the co-existence of two of the three groups, support the recognition of these groups as distinct species. The three taxa recognized at the species level are: A. undulatum Dours, 1873, which has a wide distribution in the Mediterranean and extends eastwards to Iran; A. wahrmani Mavromoustakis, 1948 stat. nov., which is found in the southern Levant; and A. libanicum Kasparek sp. nov., which is known only from Lebanon, and where it seems to be endemic. The results confirm the high diversity of anthidiine bees in the eastern Mediterranean and indicate that the species richness may be much higher than currently known. The sympatric occurrence of genetically and morphometrically distinct forms within the same habitat is considered strong evidence that these are distinct cryptic species, rather than variations arising from local geographic or environmental adaptations.

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DOI: 10.3897/asp.83.e130043

Authors: Jéssica Meneghetti, Alejandro Jose Biganzoli-Rangel, Gervásio Silva Carvalho, Andressa Paladini

Abstract: Mahanarva (Ipiranga) Fennah, 1968, currently comprises nine described species. The subgenus is characterized by an inflated postclypeus with a convex profile, longer than wide tegmina, and male genitalia with a short, laterally compressed aedeagus with a pair of slender dorsal processes. In this study, a phylogenetic hypothesis based on morphological data is presented to test the monophyly of Mahanarva (Ipiranga) and infer the relationship between its species. We present the redescriptions of its species and the description of two new species, Mahanarva (Ipiranga) obliqua sp. nov. and Mahanarva (Ipiranga) nefasta sp. nov., along with a dichotomous key for identification. Moreover we synonymized M. (I.) vittata and M. (I.) fortunata based on morphological characters and a principal component analysis. The PCA results suggested no differences on the morphospace between these species; however, cluster analysis resulted in three groups. These groups don’t show a geographic pattern structure or stable genitalic morphological differences. For phylogenetic analyses, a data matrix with 30 taxa and 56 characters was constructed. Based on the results of phylogenetic analyses, Mahanarva (Ipiranga) is recovered as paraphyletic and grouped with Mahanarva (Mahanarva) + Kanaima.

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DOI: 10.3897/asp.83.e147854

Authors: Ivan L. F. Magalhaes, Abel Pérez-González, Facundo M. Labarque, Martín Carboni, Jörg U. Hammel, Robin Kunz, Martín J. Ramírez, Mónica M. Solórzano-Kraemer

Abstract: In our previous contribution, “Revision of recluse spiders (Araneae: Sicariidae: Loxosceles) preserved in Dominican amber and a total-evidence phylogeny of Scytodoidea reveal the first fossil Drymusidae” (Arthropod Systematics & Phylogeny 80, 2022, 541–559), Table 1 was inadvertently missing from the final published version. It is included in the present Corrigendum.

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DOI: 10.3897/asp.83.e142332

Authors: Xiaozhen Lu, Jikai Lu, Yunfei Wu, Meishu Guo, Guy Smagghe, Renhuai Dai

Abstract: Iassinae, a widely distributed group of herbivorous pest insects, is a subfamily of Cicadellidae. Previous studies on the phylogeny of Iassinae were mostly based on morphological characteristics, mitochondrial genomes, and molecular fragments (H3, 28S, and 12S), and their phylogenetic relationships were controversial. To better understand Iassinae, we analyzed the phylogenetic relationships among four genera in Iassinae with use of thousands of universal single-copy orthologs and ultraconserved elements extracted from 25 newly sequenced low-coverage whole genome data. Both marker sets provided consistent results across the maximum likelihood and coalescent-based species tree approaches. The phylogenetic results showed that the two genera Batracomorphus and Trocnadella were monophyletic groups, and Krisna a paraphyletic group. For the genus Gessius, we could not explain whether it is monophyletic or paraphyletic since only one species was involved. In this study, the phylogenetic relationship with use of universal single-copy orthologs and ultraconserved elements was stable, and all results supported that Batracomorphus is a sister group of Trocnadella, and that Gessius and Krisna possess a sister relationship. In addition, the divergence time showed that the divergence of Batracomorphus, Trocnadella, Krisna and Gessius began at approximately 49–72 Mya, 33–57 Mya, 51–78 Mya and 17–36 Mya, respectively. These results will help us to understand the phylogeny and evolutionary relationship of Iassinae.

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DOI: 10.3897/asp.83.e116160

Authors: Dieter Weber, Martin Wendt, Thomas Schmitt

Abstract: The biogeography, taxonomy and systematics of ground water organisms is still poorly understood. This is partly due to the difficult accessibility of the habitats and the expert knowledge required for identification. Nevertheless, due to the large distribution range and limited dispersal possibilities of amphipods such as the genus Niphargus, important insights can be gained into biogeographical patterns and evolutionary processes in subterranean ecosystems. Niphargus is the most species-rich genus of freshwater amphipods worldwide and holds great potential for cryptic species whose identification is important for the reconstruction of biogeographic patterns and events. Therefore, we assessed the genetic patterns of Niphargus fontanus Spence Bate, 1859 (Amphipoda: Niphargidae). We sampled the species all over its current distribution and sequenced one mitochondrial and three nuclear gene fragments. We discovered that most records from France were probably misidentifications, and that the species does not occur in central and southern France. Nevertheless, the distribution area extends from Wales to Bavaria, which could make it one of the largest distribution areas within the genus. Compared with other Niphargus, the genetic diversity and differentiation of N. fontanus is low and most likely evolved since the mid-Pleistocene Transition, but reflects a clear phylogeographic pattern with about 13 genetic lineages. These apparently stand for a number of extra-Mediterranean glacial refugia from which postglacial expansion was low to moderate. However, few cases of disjunction within these genetic lineages exist, most likely resulting from rapid expansions along river Rhine which otherwise mostly acted as a dispersal barrier.

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DOI: 10.3897/asp.83.e140421

Authors: Wei Xu, Gaoji Zhang, Tangjun Xu, Ke He, Jiachen Wang, Zeyang Liu, Hongyi Liu

Abstract: Scorpions, a diverse group of arachnids consisting of over 2,000 valid species, have received limited research attention in terms of their complete mitochondrial genomes (mitogenomes). To increase the taxonomic sampling frequency of species available for study based on mitogenomes, we reconstructed the complete mitogenomes of five scorpions, Androctonus amoreuxi (Audouin, 1826), Hottentotta tamulus (Fabricius, 1798), Leiurus quinquestriatus (Ehrenberg, 1828), Lychas mucronatus (Fabricius, 1798), and Teruelius flavopiceus (Kraepelin, 1900) within the family Buthidae. These five mitogenomes had a typical circular structure, with total sizes ranging from 14,504 to 15,083 bp. Nucleotide composition analysis indicated that the sequences were biased toward A and T. The Ka/Ks ratios within 13 protein-coding genes (PCGs) were lower than 1, suggesting that they had been subject to purifying selection in Buthidae. Our analyses provide additional evidence on that, in scorpions, the majority of mitogenome rearrangements occurred in tRNAs. Moreover, the genes tRNA-Asp, tRNA-Gln and tRNA-Ile were the hotspots of rearrangement in this order. Phylogenetic analyses based on PCGs supported taxonomic relationships in this taxon. Our results might provide useful insights into the gene arrangement features of scorpion mitogenomes and lay the foundation for further studies on the family Buthidae.

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DOI: 10.3897/asp.83.e135636

Authors: Yu-Jiao Guo, Shuang-Qi An, Zhen-Gui Fang, Shu-Fei Wei, Zi-Hao Shen, Zhao-Yang Chen, Zhu-Qing He

Abstract: We recently published the revision of Goniogryllus. However, the affiliation between Qingryllus striofemorus Chen & Zheng, 1995 and Goniogryllus ovalatus Chen & Zheng, 1996 is wrong. Q. striofemorus should be transferred to the genus Goniogryllus, as Goniogryllus striofemorus comb. nov. G. ovalatus syn. nov. and Q. jiguanshanensis Liu, Zhang & Shi, 2017 syn. nov. should be junior synonyms of G. striofemorus. Therefore, the corresponding description should be changed. We thank Dr. Holger Braun for pointing out the error.

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DOI: 10.3897/asp.82.e132546

Authors: Carmen Duque-Amado, Mario García-París, Alberto Sánchez-Vialas

Abstract: Phylogeography stands as a key tool to explore evolutionary patterns and processes, playing a crucial role in delimiting evolutionary units. Identifying these units is essential for providing robust taxonomic decisions. In this study, we present a comprehensive phylogeographical framework of Scaurus uncinus (Forster, 1771) and Scaurus gigas Waltl, 1835 across the Iberian Peninsula and North-western Africa, where they are widely co-distributed, coexisting in several localities on both sides of Strait of Gibraltar. Our results show that the Strait of Gibraltar did not act as a geographical barrier for these species, revealing shared mitochondrial haplotypes and nuclear alleles between populations on both sides. However, the Souss Valley in Morocco appears to have historically served as a significant geographical barrier within the S. gigas lineage, leading to the divergence of two morphologically distinct sublineages, one to the north (S. gigas) and the other to the south (S. ferreri stat. nov.). In addition, we point out a case of cytonuclear discordance between S. uncinus and S. gigas in the southwest corner of the Iberian Peninsula, suggesting the occurrence of an ancient event of genetic introgression between the two species.

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DOI: 10.3897/asp.82.e116205

Authors: Valentine Mewis, Peter Neu, Thomas Schmitt

Abstract: The understanding of cryptic species complexes with their often highly interesting biogeographical patterns is still a crucial aspect in evolutionary biology and related disciplines. Trichoptera are a group of insects particularly rich in unresolved groups. One example is the Wormaldia occipitalis species complex in which morphological studies suggest remarkable patterns of differentiation. In order to determine genetic differentiation and phylogenetic structure, one mitochondrial (COI) and two nuclear markers (CAD, wingless) were analysed for the W. occipitalis species complex around the Alps and northwards to Germany. The morphology-defined differentiation pattern was also observed at the genetic level. The morphologically well distinguishable groups W. occipitalis and W. subterranea were identified as two genetically distant monophyletic groups with about 10 % genetic divergence of the mitochondrial marker. These two taxa likely split during the Mio-Pliocene transition. Genetic analyses revealed four subgroups within W. occipitalis and three within W. subterranea. Several possible postglacial dispersal and differentiation processes are proposed. Thereby, W. occipitalis from the western Alps and individuals of W. subterranea from the eastern Alps spread towards Central Europe after the Last Glacial Maximum. Today, both species groups are sympatric and partly syntopic in the recolonised area in western Germany but apparently allopatric in their centres of origin around the Alps. The high genetic differentiation, lack of detectable genetic evidence for hybridisation, their syntopic distribution and the morphological distinctness indicate that W. occipitalis and W. subterranea are two distinct species. The genetically determined subgroups might represent subspecies.

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DOI: 10.3897/asp.82.e136751

Authors: Rong Chen, Yi Zhao, Sergei Golovatch, Wei-Xin Liu

Abstract: A taxonomic study of the genus Skleroprotopus Attems, 1901 from nine provinces in China was conducted utilizing morphological comparisons and molecular phylogenetic analyses. The results reveal thirteen new species, i.e. Skleroprotopus yutiantianae sp. nov., S. tiankeng sp. nov., S. megistus sp. nov., S. penglai sp. nov., S. longissimus sp. nov., S. genjudi sp. nov., S. laiyuanensis sp. nov., S. longiflagellatus sp. nov., S. change sp. nov., S. ampullaceus sp. nov., S. incisodentatus sp. nov., S. multistriatus sp. nov., and S. conicus sp. nov. This significantly enriches the diversity of Skleroprotopus in China, bringing it to a total of 18 species. With the exception of S. yutiantianae sp. nov., all these species were collected in caves. In terms of their degree of adaptation to the cave environment, the latter six species are presumed troglophiles, while the others are likely troglobites. DNA-barcoding based on the COI mitochondrial gene is documented for the first time in this genus. The specific p-distances between Skleroprotopus species range from 6.6–17.0%, while intraspecific p-distances are only 0.2–1.4%. Additionally, the morphological features of male leg-pair 1, the penis and leg-pair 7 are also discussed.

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DOI: 10.3897/asp.82.e106391

Authors: Chong Li, Liang Tang

Abstract: Staphylininae, a highly diverse subfamily of rove beetles (Coleoptera: Staphylinidae), has been the subject of numerous phylogenetic studies primarily based on molecular genetics and adult morphology. This work represents an initiation of phylogenetic studies using complete immature morphology, encompassing eggs, larvae and pupae, for 27 genera of Staphylininae and two outgroups. Our findings indicate that the combination of data from all three immature stages is more phylogenetically informative than the larval data alone. The resulting maximum parsimony tree partially aligns with previous research, although certain tribal-level issues remain unresolved. Through morphological comparisons, we revealed the morphological diversity of protibia, paratergites and parasternites of abdominal segment I as examples of parallel and mosaic evolution within Staphylininae larvae. We conducted detailed character analyses to provide explanations for these phenomena. Furthermore, this study provides the first morphological data for several species of Staphylinini. Notably, we present a comprehensive study of the morphology of immature stages of Algon sphaericollis Schillhammer, 2006, the first species of the recently established subtribe Algonina with known larva. Additionally, we provide the larval morphology data for six other species: Eucibdelus sp., Platydracus pseudopaganus pseudopatricius (Müller, 1926), Platydracus marmorellus (Fauvel, 1895), Saniderus cooteri Rougemont, 2015, Saniderus sp., and Philonthus spinipes Sharp, 1874.

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DOI: 10.3897/asp.82.e131738

Authors: Vivien Cosandey, Ondřej Konvička, Emmanuel F. A. Toussaint

Abstract: The superfamily Tenebrionoidea is one of the most challenging clades in the beetle tree-of-life owing to its vast species richness and complex taxonomic history. Within this group, the family Melandryidae has long been overlooked and its systematics remains poorly known. Using available sequence data, we infer the most comprehensive phylogeny of Melandryidae to date. Our results support the polyphyly of Melandryidae with three independent clades spread across Tenebrionoidea. To accommodate these results, we restrict the status of Melandryidae and resurrect the family Osphyidae stat. rev. The third clade corresponding to the tribe Serropalpini pro parte is placed as incertae sedis within Tenebrionoidea pending further investigation and additional taxon sampling to resolve its phylogenetic placement.

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DOI: 10.3897/asp.82.e112684

Authors: Carmelo Andújar, Peter Hlaváč, Vasily V. Grebennikov

Abstract: The tribe Raymondionymini has long been neglected in phylogenetic studies. The tribe is characterized by uncertain monophyly, fluctuating taxonomic status, and a composition prone to instability. All raymondionymine weevils are wingless and have eyes either completely absent or, rarely, consisting of a single ommatidium. With body lengths predominantly below three millimeters, they inhabit deep soil environments and are infrequently collected. The core of this tribe comprises nine genera distributed in Europe and around the Mediterranean region and encompassing 76 species, while six additional genera include 17 species distributed in USA (California), Mexico, Ecuador, Venezuela, Russian Far East, and Madagascar. Here, we present eight new mitogenomes, complemented by one publicly available, encompassing all but two Mediterranean genera of raymondionymine weevils. We used publicly available Curculionoidea mitogenomes to compile an all-inclusive dataset with 391 terminals and a reduced dataset with 61 terminals representing main families of Curculionoidea and subfamilies within Curculionidae. Our maximum likelihood and Bayesian phylogenetic analyses, employing both DNA and amino acids datasets under alternative partition schemes, consistently produced congruent phylogenies. Our results show that the Mediterranean raymondionymines form a strongly supported clade, and their easternmost and morphologically distinct genus Ubychia is sister to the rest of them. Most notably, our results consistently recover a sister relationship between the clade of Mediterranean raymondionymine weevils and a clade encompassing all remaining Curculionidae. Consequently, we propose a revision of weevil taxonomy: (i) Our target group is removed from the non-monophyletic subfamily Brachycerinae; (ii) this clade is resurrected to its former subfamily level within Curculionidae, as the subfamily Raymondionyminae stat. rev; (iii) the nine Mediterranean genera Alaocephala, Alaocyba, Coiffaitiella, Derosasius, Ferreria, Raymondiellus, Raymondionymus, Tarattostichus, and Ubychia compose Raymondionyminae stat. rev; (iv) and non-Mediterranean genera Alaocybites, Bordoniola, Gilbertiola, Homosomus, Neoubychia, and Schizomicrus are considered as “incertae sedis” pending further phylogenetic corroboration. We hypothesize that the remaining Brachycerinae and the non-Mediterranean representatives within Raymondionyminae constitute a series of species-poor early-diverging lineages representing currently unrecognized subfamilies of Curculionidae.

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DOI: 10.3897/asp.82.e114351

Authors: Mengjiao Ren, Chunpo Tian, Alexandra A. Grossi, Fasheng Zou, Daniel R. Gustafsson

Abstract: Closely related chewing lice in the Philopterus-complex are typically morphologically homogeneous, with the most significant differences often being in the male genitalia. However, in many groups within this complex the male genitalia are reduced and lacking one or more element, with the remaining components often at least partially fused. This is not least the case in the genus Philopteroides Mey, 2004, in which the male mesosome is often reduced and other characters are largely homogeneous throughout the genus. A phylogeny of the group based on a combination of mitochondrial and nuclear genes suggests that the species presently placed in Philopteroides belong to three different clades, which do not form a monophyletic group together. We here redefine Philopteroides morphologically, and describe two of these clades as new genera: Stasiasticopterus n. gen. for the species on bulbuls, and Coronedax n. gen. for species on monarch flycatchers. These genera can be separated from each other and from Philopteroides only by some characters of the male genitalia, but possibly also by characters of the preantennal head and female genitalia. In addition, we describe a new species of Coronedax, Coronedax longiceps sp. n. and provide an overview of the morphology of the male genitalia in the three genera treated.

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DOI: 10.3897/asp.82.e108699

Authors: Yu-Jiao Guo, Shuang-Qi An, Zhen-Gui Fang, Shu-Fei Wei, Zi-Hao Shen, Zhao-Yang Chen, Zhu-Qing He

Abstract: Species belonging to the genus Goniogryllus Chopard, 1936 are primarily identified based on their morphological characteristics. However, male genitalia provide limited features for classification. In this study, we sequenced the COI gene from 49 samples, and reconstructed a phylogenetic tree. Species were classified by a combination of molecular and morphological data, and some morphological features were re-evaluated. The results indicate that the presence or absence of pubescence on the thorax and abdomen, yellow stripes on the head with or without bifurcation, and the color of the maxillary palpus are interspecific differences. The classification based on these morphological features are consistent with the classification based on COI gene. In contrast, variations in the color of the hind femur, pronotum, and the number of dorsal spurs on the hind tibia are considered to be intraspecific variations. Additionally, Qingryllus striofemorus Chen & Zheng, 1995 and Q. jiguanshanensis Liu, Zhang & Shi, 2017 are recognized as the long-wing forms of G. ovalatus Chen & Zheng, 1996. Therefore, Qingryllus Chen & Zheng, 1995 syn. is a junior synonym of Goniogryllus. Q. striofemorus syn. nov. and Q. jiguanshanensis syn. nov. are junior synonyms of G. ovalatus. Furthermore, G. cirilinearis Xie, 2005 syn. nov. is a junior synonym of G. atripalpulus Chen & Zheng, 1996. G. octospinatus Chen & Zheng, 1995 syn. nov. is a junior synonym of G. chongqingensis Chen & Zheng, 1995. Lastly, Callogryllus yunnanus Wu & Zheng, 1992 is transferred to the genus Goniogryllus, as Goniogryllus yunnanus comb. nov.

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DOI: 10.3897/asp.82.e114286

Authors: Pedro Bonfá-Neto, Frederico Falcão Salles, Albane Vilarino

Abstract: Asymmetrical genitalia are reported from major Trichoptera subgroups and evolved multiple times independently. In Oecetis, it is a characteristic of the insconpicua group. However, certain species in other species groups also evolved an asymmetrical spiny projection on the phallotheca. Here, two new species with an asymmetric projection are described in the falicia group from the Brazilian Atlantic Forest, Espírito Santo state. Their phylogenetic relationships were investigated through a Bayesian analysis combining COI and morphological data. Additionally, we provide new records of Oecetis connata, O. inconspicua and O. paranensis from the Espírito Santo state, and O. connata and O. flinti from Minas Gerais state. Oecetis capixaba sp. nov. is placed with low support as the sister species of O. acanthostema, both presenting stout spine-like setae on the inner surface of the inferior appendage; the new species is diagnosed by the long spine-like setae on the inferior appendage, the narrow dorsolateral process of segment IX, and the phallic apparatus without apical projections. Oecetis ruschii sp. nov. is placed as a sister group of the clade including O. facilia and O. furcata, both presenting forked dorsolateral processes of segment IX; the new species is diagnosed by the dorsolateral process of segment IX with a lateral branching and the apex of inferior appendage wide and triangular. While the phylogenetic results should be considered preliminary and interpreted with caution, they indicate that the asymmetric projection evolved multiple times convergently in the avara, punctata, and falicia groups. The asymmetric genitalia in the falicia group seems to have evolved only in males since no correspondent asymmetry is described for females. The function of the asymmetrical projection remains unknown.

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DOI: 10.3897/asp.82.e113041

Authors: Jin Wang, Chong-Hui Yao, Chao Jiang, Wei-Chun Li

Abstract: The genus Lucasioides Kwon, 1993 (Isopoda: Oniscidea: Agnaridae) occurs in Asia, but confined only to China, Japan, Korea, and Russian Siberia. The ambiguously morphological differences among some members of the genus make a dilemma: the species recognition, whether morphologically similar or different, is uncertain. In this paper, we present first morphometric and molecular data for the genus from a broad sample. DNA sequences (mitochondrial COI, nuclear 18S rRNA, 28S rRNA, and NaK) were generated and integrated with morphological evidence to reveal the cryptic species and delimit the new species within the genus. Seven species are described as new to science: Lucasioides digitatus sp. nov., L. dissectus sp. nov., L. dianensis sp. nov., L. formosus sp. nov., L. gracilentus sp. nov., L. schmidti sp. nov. and L. subcurvatus sp. nov. To date, Lucasioides species from China are 44.4% as many as all the known congeners worldwide, showing the exceptional species diversity of Lucasioides species from China. The results demonstrate that the integrative taxonomy is especially important to reveal the cryptic species among the high morphological similarity of taxa, as well as providing an effective way for species identification to accelerate the exploration of woodlice biodiversity.

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DOI: 10.3897/asp.82.e115954

Authors: Hoi-Sen Yong, Sze-Looi Song, Kah-Ooi Chua, Yvonne Jing Mei Liew, Kok-Gan Chan, Phaik-Eem Lim, Praphathip Eamsobhana

Abstract: Bactrocera (Bulladacus) cinnabaria and B. (Bactrocera) propinqua are tephritid fruit flies of the subfamily Dacinae, tribe Dacini. The whole mitogenomes of these two species (first report for the subgenus Bulladacus) possess 37 genes (13 protein-coding genes – PCGs, 2 rRNA and 22 tRNA genes). The mitogenome of B. cinnabaria (15,225 bp) is shorter than that of B. propinqua (15,927 bp), mainly due to the smaller size of the control region and intergenic spacers in B. cinnabaria. Molecular phylogeny based on mitochondrial genes (mt-genes) reveals two clades of the genus Bactrocera: one comprising the subgenus Bactrocera and the other comprising the subgenera Bulladacus, Daculus, Tetradacus and unassigned Bactrocera sp. ‘yunnanensis’. The subgenera represented by two or more taxa are monophyletic. B. (Bulladacus) cinnabaria forms a sister group with the subgenus Tetradacus (B. minax and B. tsuneonis) and B. sp. ‘yunnanensis’, in a clade containing also the basal sister lineage of the subgenus Daculus (B. oleae and B. biguttula). B. propinqua forms a sister group with B. ritsemai and B. limbifera in a subclade containing also B. umbrosa, B. curvifera and B. moluccensis of the monophyletic subgenus Bactrocera. The present study supports the synonymy of B. ruiliensis with B. thailandica. It also shows a high genetic similarity between (a) B. melastomatos and B. rubigina, (b) B. papayae and B. philippinensis, (c) B. dorsalis and B. invadens, (d) B. tryoni and B. neohumeralis, and (e) B. cheni and B. tuberculata; and B. cheni is distinct from and not a synonym of B. tsuneonis or B. lombokensis.

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DOI: 10.3897/asp.82.e116541

Authors: Zoltán Kenyeres, Norbert Bauer, Maciej Kociński, Beata Grzywacz

Abstract: Steppes form large zonal habitats in Asia but only consist of localised outposts in Europe. An ideal subject for the study of differences within species between the main steppe zone and the localized more western outposts is the Orthopteran Stenobothrus eurasius, widespread across the Siberian and Central Asian steppes but present only in isolated relic populations at the western edge of its area. We used genetic and morphological analyses to detect possible differences among these relic populations. We carried out a study on morphological parameters of wings in parallel with the comparison of four DNA fragments (cytochrome c oxidase subunit I, 12S rRNA and the mtDNA control region, cytochrome B, nuclear internal transcribed spacers plus the 5.8S rRNA region) involving 15 extrazonal populations of the species. St. nigromaculatus was used as an outgroup taxon in the genetic analyses. Variability of the morphological characters of St. eurasius individuals was higher within the regions than amongst the regions. The two Stenobothrus species were not separated based on the CR gene. Samples of both Stenobothrus species were separated on the COI, cytB and ITS1-5.8S-ITS2 phylogenetic trees with high support (PP = 1) in Bayesian analyses but clear genetic lineages were not revealed, and populations of the focal species were not grouped according to their geographic locations. The similarity of this species in different steppe outposts supports the hypothesis that St. eurasius was widespread in the more extensive steppe areas that were once present, but the extension of agricultural landuse reduced the steppe habitats resulting in the current patchy distribution of St. eurasius limited to the remaining habitats.

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DOI: 10.3897/asp.82.e113140

Authors: Noémie M. C. Hévin, Gael J. Kergoat, Alberto Zilli, Claire Capdevielle-Dulac, Boaz K. Musyoka, Michel Sezonlin, Desmond Conlong, Johnnie Van Den Berg, Rose Ndemah, Philippe Le Gall, Domingos Cugala, Casper Nyamukondiwa, Beatrice Pallangyo, Mohamedi Njaku, Muluken Goftishu, Yoseph Assefa, Onésime Mubenga Kandonda, Grégoire Bani, Richard Molo, Gilson Chipabika, George Ong’amo, Anne-Laure Clamens, Jérôme Barbut, Bruno Le Ru

Abstract: In this study, we reassess the phylogenetic relationships of the genus Sesamia Guenée, 1852 and examine in more detail the members of the nonagrioides species group, for which three distinct species complexes are identified. The calamistis subgroup comprises eight species, of which four new species are described: Sesamia kabirara Le Ru sp. nov., Sesamia kalale Le Ru sp. nov., Sesamia mapalense Le Ru sp. nov. and Sesamia teke Le Ru sp. nov. The incerta subgroup consists of 11 species, of which four new species are described: Sesamia kamba Le Ru sp. nov., Sesamia lalaci Le Ru sp. nov., Sesamia lusese Le Ru sp. nov. and Sesamia msowero Le Ru sp. nov. The nonagrioides subgroup comprises ten species of which two new species are described: Sesamia libode Le Ru sp. nov. and Sesamia satauensis Le Ru sp. nov. Phylogenetic and molecular species delimitation analyses of a multi-marker molecular dataset allow us to investigate and clarify the status of Sesamia species and species complexes. Our results yield a well-supported phylogenetic hypothesis for the genus, which supports the monophyletic nature of all but one species subgroup. The results of 16 distinct molecular species delimitation analyses show some levels of incongruence and, overall, a tendency towards over-splitting. We also present an updated list of species for the genus Sesamia and provide morphological keys based on male and female genitalia to determine the species group of any Sesamia species and to identify all species belonging to the nonagrioides species group.

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DOI: 10.3897/asp.82.e114349

Authors: Feiyang Liang, Xingyue Liu

Abstract: Psocomorpha, as one of the suborders of the order Psocodea, represents a large group of free-living bark lice. The phylogenetic relationships among/within the major groups, such as families, subfamilies, tribes, etc., of the bark lice need extensive investigation. The family Stenopsocidae belongs to the infraorder Caeciliusetae, which is a large group of Psocomorpha and is one of the common groups in Asia. Here we infer the intergeneric relationships of Stenopsocidae based on the mitochondrial genes and the nuclear 18S rRNA. The result supports the monophyly of Stenopsocidae and suggests the paraphyly of the genus Stenopsocus. Combining the morphological evidence, we propose a new genus, namely Neostenopsocus gen. n., which includes a number of species previously placed in Stenopsocus. The checklist of this new genus is provided. This new genus can be distinguished from Stenopsocus by the glabrous forewing CuP. In the revised classification system, Stenopsocidae includes four genera: Graphopsocus, Malostenopsocus, Neostenopsocus gen. n., and Stenopsocus. A key to the genera of Stenopsocidae is provided. We also present the diagnostic characters for each of these genera.

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DOI: 10.3897/asp.82.e112587

Authors: Albane Vilarino, Adolfo R. Calor

Abstract: Geometric morphometric statistics have been employed to reduce the subjectivity of visual evaluations in taxonomy. Taxonomy in most insect groups relies strongly on male genitalia morphology which is often the structure with most data available, which is also true to caddisfly taxonomy. Here we revise the caddisfly subgenus Xiphocentron (Sphagocentron) adding five new species after 40 years: X. dactylum sp. nov., X. eurybrachium sp. nov., X. tapanti sp. nov., and X. tuxtla sp. nov. Additionally, we describe a new X. (Antillotrichia): X. drepanum sp. nov. from French Guiana and provide new species records of Xiphocentronidae from Bolivia, Costa Rica and Ecuador. We performed exploratory geometric morphometric analysis on the male genitalia’s preanal appendage to characterize the shape differences among the species, and to investigate its utility to classify species to subgenera. In order to infer species relationship and assess if shape congruences are due to phylogenetic signal or convergence data from 100 landmarks and semilandmarks, and 30 discrete characters were used to generate a phylogenetic hypothesis. The morphometry partially supports the subgenera delimitations, but the Antillotrichia subgenus greatly overlapped with other subgenera. The discriminant analysis overall classification correctness was 64%. Some suggested phenotypic groups were due to convergence. According to the preanal appendage morphometry, X. (Antillotrichia) fuscum is a Sphagocentron species. The phylogenetic analysis recovered Sphagocentron as monophyletic, but not Antillotrichia. Sphagocentron subgenus was placed within a clade of several Antillotrichia species, with X. (A.) fuscum as the sister of the other Sphagocentron species, although support values were low.

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DOI: 10.3897/asp.82.e115268

Authors: Nielson Felix Caetano França, Célio Magalhães, Fernando Luis Mantelatto

Abstract: The taxonomic status of the freshwater crab Dilocarcinus septemdentatus (Herbst, 1783) is still not well established. Currently, the main issue involves synonymization with D. spinifer H. Milne Edwards, 1853, based on a variation of the angulation of the gonopod apex. These species are distributed along rivers and lakes in northern South America, with disjunct occurrences in central-west Brazil and Argentina. Due to these inconsistencies, an integrative approach was performed to elucidate these questions, with morphological (including NanoCT-Scan) and molecular analysis (Maximum Likelihood Trees, Bayesian Inference, Genetics Distance Matrix, and Haplotype Network), based on mitochondrial markers COI and 16S rRNA. Both analysis revealed and supported the existence of a species complex under the name of D. septemdentatus. Based on the results obtained, we propose the revalidation of D. spinifer, the description of a new species, and the redescription of D. septemdentatus s. str., with a neotype designation for this species. The hypothesis that this species complex originated in the Pebas System, an extensive mega wetland system that existed along the lowlands of Western Amazonia from late Oligocene to late Miocene (c. 23–11 mya) is discussed.

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DOI: 10.3897/asp.82.e114213

Authors: Jamie Ramirez, Agnieszka Bugaj-Nawrocka, Artur Taszakowski, Christiane Weirauch

Abstract: The assassin bug genus Koenigsbergia Popov, 2003 is currently monotypic and represented by a female holotype from Baltic Amber (~33.9–55.8 MYA). The genus was originally described within Phymatinae (Phymatine Complex or phymatine clade). However, our literature review reveals that the amber fossil likely belongs to the subfamily Phimophorinae, which is distantly related to the phymatine clade. The recent acquisition of one male and one nymph of Koenigsbergia provides the opportunity to reevaluate the systematic placement of this genus. We here examine the new fossils, concluding that the adult male represents an undescribed species, and describe it as Koenigsbergia explicativa, new species. Our morphological comparison between Phimophorinae, Phymatinae, and Koenigsbergia (macro imagining, scanning electron microscopy) shows that the fossil genus shares notable similarities with Phimophorus Bergroth, 1886 and Mendanocoris Miller, 1956. We therefore formally transfer the fossil genus to Phimophorinae.

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DOI: 10.3897/asp.82.e114038

Authors: Adrian Smolis, Nataliya Kuznetsova, Grzegorz Paśnik

Abstract: With 54 described species, Endonura is one of the most species-rich genera of the Neanurinae. The taxon is characterised by the presence of 0–2 ocelli, separate tubercles Di and De with the non-crossed type of chaetotaxy on the head, and two or three tubercles on the fifth abdominal tergite. Four new species from the Crimea and the Caucasian region have been described based on adult specimens: Endonura armeniaca sp. nov., E. cochlearifera sp. nov., E. crimica sp. nov., and E. duplex sp. nov. A phylogenetic analysis of the genus Endonura using 70 characters of adult external morphology is presented. Two methods were chosen to examine different approaches to the reconstruction of evolutionary relationships: Maximum Parsimony (MP) and Bayesian Inference (BI). Besides supporting Endonura monophyly both analyses failed to resolve any of the deeper relationships within the genus. All known species within the genus are grouped into two main clades, A and D. Both of these clades have members distributed throughout Europe, the Caucasus and the western part of Central Asia. Endonura crimica sp. nov. can be considered as an intermediate between these two clades.

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DOI: 10.3897/asp.82.e98662

Authors: Masoumeh Amiri, Lorenzo Prendini, Fenik Sherzad Hussen, Mansour Aliabadian, Roohollah Siahsarvie, Omid Mirshamsi

Abstract: Morphological and genetic variation among populations of the widespread buthid scorpion, Hottentotta saulcyi (Simon, 1880), occurring in western and southwestern Iran was explored using morphometric variables, one nuclear marker (28S rDNA) and three mitochondrial markers (12S rDNA, 16S rDNA, and Cytochrome c Oxidase Subunit I). Genetic and morphometric statistical analyses revealed extensive cryptic diversity. Phylogenetic analysis with Bayesian Inference and Maximum Likelihood uncovered two divergent clades, one of which is described as a new species, Hottentotta hatamtiorum sp. nov., from Ilam and Khuzestan Provinces, southwestern Iran. The description of the new species raises the total count of Hottentotta Birula, 1908 species to 61, twelve of which are endemic or subendemic to the Iranian Plateau.

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DOI: 10.3897/asp.82.e116703

Authors: Kinga Walczak, Thomas Pape, James F. Wallman, Krzysztof Szpila, Andrzej Grzywacz

Abstract: The muscid genus Achanthiptera Rondani (Diptera: Muscidae) was classified within its own subfamily Achanthipterinae for decades due to a misinterpretation of adult morphology. Conversely, the larval morphology suggested that Achanthiptera should be classified within Azeliinae, yet no formal changes were implemented based on this source of data. Using scanning electron microscopy (SEM) and confocal laser scanning microscopy (CLSM), we examined the larval morphology of Ac. rohrelliformis (Robineau-Desvoidy), Potamia littoralis Robineau-Desvoidy and Australophyra rostrata Robineau-Desvoidy. Despite the challenges posed by the poor condition of hundred-year-old museum specimens of Ac. rohrelliformis for light microscopy, CLSM examination yielded satisfactory results. Additionally, CLSM observations revealed peculiar modifications to the cephaloskeleton, including a dome-shaped (second instar) or spade-like (third instar) anterior rod attached to each mouthhook in Ac. rohrelliformis and P. littoralis. These structural modifications are likely to enhance the efficiency of food collecting by enlarging the surface of the mouthhooks. The results of our morphological analyses lead to the conclusion that larvae of Au. rostrata are facultative carnivores, while modified accessory oral sclerites in Ac. rohrelliformis and P. littoralis suggest a saprophagous feeding strategy. This study contributes new evidence that Achanthiptera is the sister taxon to Potamia Robineau-Desvoidy, and both are nested within the subfamily Azeliinae.

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DOI: 10.3897/asp.82.e114508

Authors: Erich L. Spiessberger, Alfred F. Newton, Margaret K. Thayer, Oliver Betz

Abstract: The mouthparts and protarsi of adult rove-beetles of the Staphylinine group are examined in detail. We provide descriptions and image plates based on scanning electron micrographs taken from 36 species representing all 10 subfamilies comprising this large staphylinid subunit. We establish groundplan features of the mouthparts for the Staphylinine group and discuss, in detail, aspects and functions of structures that compose the feeding apparatus. A phylogenetic scheme is used to conduct an ancestral character reconstruction of the morphological characters. The potential groundplan features of the characters rendered in our parsimony analysis for the Staphylinine group are: labrum subquadrate or longer than wide; mandible without subapical teeth and retinaculum, with prostheca present, not forming lobe-like projection, and with a mola; maxillary palpomere 4 well-developed, fully sclerotized, similar in width to palpomere 3; ‘glossa’ integrated with prementum plate, sometimes represented by pairs of sensilla basiconica; ‘paraglossa’ with unmodified antero-lateral lobes; labial palpomere 3 from as wide to half as wide as penultimate palpomere. To explain the shape variation of the mandibles, a geometric morphometric analysis was carried out. A character mapping analysis of mandible shapes revealed a trend in the Staphylinine group toward a falcate shape with a narrow base, typically present in some predatory species.

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DOI: 10.3897/asp.82.e105740

Authors: Atefe Kalate, Alireza Keikhosravi, Reza Nasrabadi, Elahe Parvizi, Christoph D. Schubart, Reza Naderloo

Abstract: The current patterns of biodiversity have significantly been affected by glacial-interglacial cycles during the Pleistocene period. True freshwater crabs are considered as poor dispersers and terrestrial barriers restrict gene flow between their local populations. Recent studies, however, suggest that certain environmental conditions, such as periods of heavy rainfall and humidity, can facilitate their between-drainage dispersal and will result in the evolution of homogenous genetic patterns among different drainage systems. Here we tested this hypothesis by comparing populations of the endemic freshwater crab Potamon elbursi Pretzmann, 1962 distributed in two drainage systems, the Caspian Sea and Namak Lake, in northern Iran. Our results based on the genetic analysis of 70 new and 61 previously published sequences of the mitochondrial cytochrome oxidase subunit I gene revealed a substantial haplotype diversity in some populations and high levels of local population structuring in others. Initially, we found mixed evidence of genetic differentiation and connectivity among drainages and populations. Genetic differentiation between the two drainages only became apparent after the Sepirdood population (which belongs to the Caspian Sea drainage) was excluded. Subsequently, the two drainages showed significant genetic distinctions, with a limited gene flow between them. Our demographic analyses supported recent population bottlenecks, followed by a rapid demographic and/or spatial expansion dating back to the Pleistocene climatic fluctuations. Species distribution modelling suggests that precipitation during warm weather conditions profoundly affects the distribution of P. elbursi. This study indicates that freshwater crabs can override short land barriers under favorable weather conditions and will have conservation implications in the face of contemporary climatic fluctuations.

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DOI: 10.3897/asp.82.e106734

Authors: André L. Martins, Gabriel A. R. Melo

Abstract: Thaumatodryininae is a small subfamily of Dryinidae, known to attack nymphs of auchenorrhynchous Flatidae (Hemiptera). Only one genus is recognized, Thaumatodryinus Perkins, with 35 species including fossil and extant taxa. Currently, the oldest record for the genus is from Baltic amber. Here, we present the first record of Thaumatodryininae from mid-Cretaceous Burmese amber with the description of †Thaumatorrhinos athrix gen. et sp. nov., derived from the first phylogeny for this subfamily based on morphological characters. The placement of †Thaumatorrhinos gen. nov. in Thaumatodryininae and the phylogenetic relationships of this subfamily within Dryinidae are discussed.

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DOI: 10.3897/asp.82.e104232

Authors: Zhengyang Liu

Abstract: The moth genus Sinobirma was reared successfully for the first time, based on specimens of Sinobirma bouyeri collected in the southeastern Himalayas of Tibet. Larvae were reared on the host plants Coriaria nepalensis and Prunus cerasoides in captivity in Yunnan. Morphology and biology of the ovum, larvae, and pupa of S. bouyeri are described in detail. The species exhibits strong gregarious behavior during all larval instars, with mature larvae of S. bouyeri primarily feeding at night. The larvae are black and decorated with green stripes, pupating individually in the soil. Numerous host plants known to be used by African and Asian Saturniidae were tested with larvae of this species. The first parasitoid for the genus Sinobirma is reported. The complete mitochondrial genome was sequenced and used to reconstruct a molecular phylogeny to test the tribal placement of Sinobirma. The paper provided further evidence that Sinobirma originated from the African mainland and reached the Himalayas through dispersal.

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DOI: 10.3897/asp.82.e114849

Authors: Sergio Álvarez-Parra, André Nel, Vincent Perrichot, Corentin Jouault

Abstract: The order Psocodea, including barklice, booklice, and parasitic lice, is diverse and widely distributed since the Cretaceous. That is particularly the case for the speciose extinct family Empheriidae (Psocodea, Trogiomorpha, Atropetae), recently fused with the ‘Archaeatropidae’. Understanding the evolution of barklice is dependent in part on studying this family, as its representatives have been found from the Early Cretaceous to the Eocene, surviving the K/Pg extinction event. The phylogenetic relationships of Empheriidae in relation to other families, such as Lepidopsocidae or Psoquillidae, have been extensively debated. However, distinguishing diagnostic characters for the Empheriidae has proven challenging. In this study, we describe the new empheriid Santonipsocus mimeticus gen. et sp. nov. from Cretaceous Charentese amber (France). It is the third empheriid species found in this locality. The new genus is compared with the other genera in the family, and Proprionoglaris guyoti and Proprionoglaris axioperierga are revised based on the type material and new specimens. We explore the phylogeny of Empheriidae, both the relationships with other families and the inner relationships between the genera, through maximum parsimony analysis and Bayesian inference analysis. Our results suggest that Empheriidae may represent a paraphyletic evolutionary grade to the rest of Atropetae. The phylogenetic relationships between genera align with the biogeography of the family and support previous hypotheses. In addition, we discuss the possible biology of the members of the family, shedding light on the evolutionary history of Empheriidae.

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DOI: 10.3897/asp.82.e111148

Authors: Manuel Brazidec, Lars Vilhelmsen, Brendon E. Boudinot, Adrian Richter, Jörg U. Hammel, Evgeny E. Perkovsky, Yong Fan, Zhen Wang, Qiong Wu, Bo Wang, Vincent Perrichot

Abstract: The Megalyridae are a small family of parasitoid wasps comprising eight extant genera (71 species) and six extinct genera (13 species). Here, we report eight new species from Late Cretaceous Kachin (Myanmar) and Taimyr (Russia) ambers; the family is recorded for the first time from the latter. †Cretolyra noijebumensis gen. et sp. nov., †Cretolyra shawi gen. et sp. nov., †Genkyhag innebula gen. et sp. nov., †Megacoxa chandrahrasa gen. et sp. nov., †Megacoxa janzeni gen. et sp. nov., and †Megacoxa synchrotron gen. et sp. nov., are described from late Albian – early Cenomanian Kachin amber; †Kamyristi exfrigore gen. et sp. nov. and †Kamyristi yantardakhensis gen. et sp. nov. from Taimyr amber (Baikura, late Albian – early Cenomanian Ognevka Formation and Yantardakh, Santonian Kheta Formation, respectively). Phylogenetic analyses of the family are presented and the classification of the Megalyridae is revised accordingly. A new tribe, †Megalavini trib. nov., is erected to accommodate †Cretolyra gen. nov., †Genkyhag gen. nov., †Megacoxa gen. nov. together with †Megalava Perrichot, 2009; †Megalavini trib. nov. is characterized by the vein M fully pigmented and veins Rs+M and M+Cu aligned, the crenulate mesometapectal sulcus, and the pronotal spiracle not posteriorly surrounded by cuticle. This spiracular condition was previously only observed in †Megazar Perrichot, 2009, thus the subfamily †Megazarinae Perrichot, 2009 stat. nov. is proposed for the clade (†Megalavini + †Megazar), as sister to the remaining Megalyridae. The latter is defined as the Megalyrinae, with †Kamyristi gen. nov. being retrieved as sister to all other genera except Rigel Shaw, 1987 under parsimony analyses. To align phylogeny with classification, three new tribes are erected (†Kamyristini trib. nov., †Megallicini trib. nov., and †Valaaini trib. nov.) and the †Cretodinapsini is synonymized under Megalyrini. A grouping [Rigel + Megalyridia] is supported under Bayesian analyses, which is the only specific conflict with the parsimony analyses, suggesting some degree of caution with respect to the internal relationships of the Megalyrinae. Finally, a revised key to the Megalyridae genera is provided.

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DOI: 10.3897/asp.82.e114299

Authors: Yiming Niu, Fengming Shi, Xinyu Li, Sainan Zhang, Yabei Xu, Jing Tao, Meng Li, Yuxuan Zhao, Shixiang Zong

Abstract: Longhorn beetles (Cerambycidae) play a vital role in global ecosystems. Some of them contribute to nutrient cycling and pollination, while others, pose a threat to forestry production. Despite their ecological importance, there has been a lack of comprehensive analyses on the mitochondrial genomes of Cerambycidae beetles. In this study, we have conducted mitochondrial genome sequencing and annotation for four Cerambycidae beetles: Monochamus sutor, Monochamus guerryi, Monochamus galloprovincialis, and Monochamus latefasciatus. Our analysis revealed a high degree of conservation in these mitochondrial genomes, with rare gene rearrangements observed across the Cerambycidae family. Additionally, a notable bias towards AT content was identified, with most genes using ATN as the start codon and TAA as the stop codon. Except for trnS1, all tRNA genes showed typical cloverleaf secondary structures. Phylogenetic analysis using IQ-TREE and Phylobayes consistently produced congruent topologies. At the gene level analyses, our results highlighted the remarkable conservation of the COX1 gene. Furthermore, at the species level, we observed strong adaptability in the Spondylidinae and Lepturinae subfamilies. We also offer our insights into contentious aspects of the phylogeny. Overall, our research contributes to a deeper understanding of the phylogeny and evolution of Cerambycidae, laying the groundwork for future population genetic investigations.

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DOI: 10.3897/asp.82.e105560

Authors: Yuan Hua, Ning Li, Jian Su, Baozhen Hua, Shiheng Tao, Lianxi Xing

Abstract: The scorpionfly genus Furcatopanorpa Ma & Hua, 2011 is a monotypic taxon of Panorpidae with a series of unique characters. However, the phylogenetic position of Furcatopanorpa in Panorpidae has not been satisfactorily resolved yet. Based on 48 complete mitochondrial genomes, we analyzed the mitochondrial phylogenomics and phylogeny of representatives of Panorpidae. The phylogenetic analyses indicate that Furcatopanorpa and Neopanorpa form a sister group relationship with high support. The chronogram of Panorpidae shows that Furcatopanorpa and Neopanorpa separated at ca. 82.07 Ma, while the species of Neopanorpa shared the most recent common ancestor at 49.07 Ma. Judged from the topology of the phylogenetic trees, it seems unsuitable to assign Furcatopanorpa into the subfamily Panorpinae, because this assignment may cause Panorpinae to be a paraphyletic group. A putative conclusion might be that Furcatopanorpa may need to be raised to subfamily status.

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DOI: 10.3897/asp.82.e112834

Authors: Zohreh Mirzaee, Roberto Battiston, Francesco Ballarin, Saber Sadeghi, Marianna Simões, Martin Wiemers, Thomas Schmitt

Abstract: The dwarf-mantid genus Holaptilon Beier, 1964 is composed of small-sized ground-runner species distributed in the Middle East. Due to their elusive lifestyle, little is known about their behaviour, distribution, and phylogeny. The genus Holaptilon was once established for a single species, H. pusillulum Beier, 1964, based on material collected in Jerusalem, Israel. Later, H. brevipugilis Kolnegari, 2018, and H. yagmur Yılmaz and Sevgili, 2023 were described from Iran and Turkey, respectively. In this study, integrated morphology, molecular analyses, and ecology were used to revise the genus Holaptilon and define the boundaries of its species. New data on this genus are presented, based on Holaptilon specimens collected from various provinces of Iran, Israel, Jordan, and Turkey. Extensive analyses, including examinations of male and female genitalia, morphometrical analysis, and morphological hypervolumes were conducted to distinguish its species morphologically. In addition, four molecular markers (mitochondrial and nuclear) were studied to gain a better understanding of species delimitation and phylogenetic relationships. As a result, impressive inter- and intraspecific variability was recovered. In addition to the three already known species, four new species with their distributions restricted to Iran (H. abdullahii sp. nov., H. khozestani sp. nov., H. iranicum sp. nov., and H. tadovaniensis sp. nov.) are here described, and H. yagmur Yılmaz and Sevgili, 2023 is synonymized with H. brevipugilis Kolnegari, 2018. The integrative approach was essential for an adequate classification in Holaptilon taxonomy and also helpful in the clarification of problematic and cryptic Mantodea species. Additional information concerning the life cycle, ecological aspects, spermatophore feeding, as well as geographic range and historical biogeography of Holaptilon species is also provided.

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DOI: 10.3897/asp.82.e108906

Authors: Bao-Jun Xie, Ping-Ping Chen, Jakob Damgaard, Jie-Yi Xie, Qiang Xie, Yan-Hui Wang

Abstract: The genus Micronecta Kirkaldy, 1897 is the most species-rich genus in the family Micronectidae, containing more than 160 species. Micronecta is currently divided into 11 subgenera, five of which are monotypic. Moreover, the subgenus Micronecta is an empirical mixture group. The definitions of some subgenera were based on only a few aberrant morphological features, which are specializations with few phylogenetic significances. The relationship between these subgenera remains unclear. In this study, we newly sequenced mitochondrial genomes (mitogenomes) and nuclear rDNAs (nrDNAs) for 13 Micronecta species, representing seven subgenera, and those for ten other water bugs. Our phylogenetic analyses showed that the subgenus Lundbladella represents the sister group to all other studied subgenera of Micronecta. The subgenus Unguinecta was the sister group to the clade that contains Dichaetonecta and Sigmonecta. More importantly, the subgenus Micronecta represents a paraphyletic group, which further forms a monophyletic group together with the subgenera Basileonecta and Ctenonecta. This is for the first time that the phylogeny of the genus Micronecta was investigated based on molecular data and the paraphyly of the subgenus Micronecta was revealed. Such evidence suggested the necessity of the revision of the taxonomic system of the genus in the future, and may also serve as a reference for the delimitation of subgeneric characters.

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DOI: 10.3897/asp.82.e96452

Authors: Shota Shimizu, Ryuichiro Machida

Abstract: Abstract We examine and describe the embryonic development of 13 species from eight families of Dermaptera, i.e., all families excluding Karschiellidae, Hemimeridae, and Arixeniidae: Diplatys flavicollis (Diplatyidae), Cranopygia sp., Echinosoma sp., and Parapsalis infernalis (Pygidicranidae), Apachyus chartaceus (Apachyidae), Anisolabis maritima and Euborellia pallipes (Anisolabididae), Labidura riparia (Labiduridae), Forficula scudderi and Anechura harmandi (Forficulidae), Paralabella curvicauda (Spongiphoridae), and Proreus simulans and chelisochid gen. sp. (Chelisochidae). We also provide new findings on the reproductive biology of the Pygidicranidae and the postembryonic development of the Apachyidae. Based on information from the present and previous studies, we reconstruct the developmental and reproductive-biological groundplan for Dermaptera and discuss phylogenetic issues related to this order. We confirmed that Dermaptera possesses the embryological features (related to mode of embryonic formation and manner of blastokinesis) that are regarded as autapomorphies of Polyneoptera. Eudermaptera is characterized by the extraordinarily great length of the embryo which attains its maximum length in anatrepsis period, the positioning of its posterior end at the egg’s anterior ventral side, the type of egg tooth, and four larval instars. Anisolabididae, Labiduridae, and Eudermaptera share an elongation ratio of embryos in the anatrepsis period (ERE) of 160% or less and a larval instar number of five or less, whereas Protodermaptera is characterized by an ERE of 210% or more, a ratio of embryonic primordium relative to the egg’s longitudinal circumference (IL) of 40% or less, and a larval instar number of six or more. Notably, the ERE, IL, and larval instar number of Apachyidae are within the ranges observed in Protodermaptera.

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DOI: 10.3897/asp.82.e111985

Authors: Carol-Anne Villeneuve, Louwrens P. Snyman, Emily J. Jenkins, Nicolas Lecomte, Isabelle Dusfour, Patrick A. Leighton

Abstract: Abstract Arctic ecosystems face increasing risks from vector-borne diseases due to climate-driven shifts in disease patterns and vector distribution. However, species identification challenges impact vector-borne disease surveillance, necessitates accurate identification. Aedes species are predominant among Arctic mosquitoes and pose health risks, with some species potentially carrying Jamestown Canyon and Snowshoe hare viruses. However, identifying Aedes species is challenging, especially under Arctic conditions and with complex adult traits. This study assessed the suitability of DNA barcoding (COI and ITS2 regions) and morphological characteristics for the identification of Arctic black-legged Aedes. It also aimed to evaluate the reliability of publicly available sequences. Our analysis focused on Aedes impiger, Aedes nigripes, and two species from the Punctor subgroup – Aedes hexodontus and Aedes punctor. In our study, the COI barcoding region distinguished Ae. impiger and Ae. nigripes but not within the species of the Punctor subgroup. In addition, the ITS2 barcoding region did not differentiate the species. When we evaluated GenBank and BOLD sequences, we found issues of under-representation and misidentifications, particularly within the Punctor subgroup. Based on these results, we recommend addressing identification difficulties, particularly within the Punctor subgroup, and advocate for more comprehensive morphological and molecular identification strategies. Integrating morphology and DNA barcoding holds promise for robust disease surveillance in Arctic regions, yet challenges persist, especially in complex species groups like the Punctor subgroup. Tackling these issues is pivotal to ensuring accurate vector status determination and reliable disease risk assessments in a rapidly changing Arctic ecosystem.

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DOI: 10.3897/asp.82.e109995

Authors: Daubian Santos, Guilherme Cunha Ribeiro

Abstract: Abstract Eloeophila Rondani, 1856 and Idioptera Macquart, 1834 are two genera of the family Limoniidae (Diptera) distinguished by the presence of a supernumerary cross-vein m-cu. Although these genera were previously combined, there has been a lack of phylogenetic tests to investigate the evolutionary relationship between them. In this study, we conducted a cladistic analysis that indicates that Idioptera form a clade within Eloeophila, and therefore the two genera should be synonymized under Idioptera. Consequently, 87 species of Eloeophila are transferred to Idioptera.

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DOI: 10.3897/asp.81.e111799

Authors: Hossein Ashrafi, Kristin M. Hultgren

Abstract: Abstract The alpheid snapping shrimp genus Synalpheus Spence Bate, 1888, is a prominent component of arthropod diversity found in coral reefs. Notably, Synalpheus is the only genus of marine organisms known to exhibit eusocial behavior. Although eusociality has evolved at least four times independently in Synalpheus, all described eusocial species are from the West Atlantic, with only a single study documenting possibly eusocial species from Indonesia. In 2008, during an expedition to Madagascar organized by the Florida Museum of Natural History (FLMNH), a diverse array of sponge-dwelling species was collected, including two species of Synalpheus exhibiting colonial behavior. Through detailed examination of these specimens, we have confirmed that these two species are eusocial and represent new eusocial species of Synalpheus outside of the West Atlantic. Consequently, we provide the first official documentation of eusocial species from the Western Indian Ocean and present their taxonomic descriptions and their phylogenetic relationships with other species of the genus in this study.

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DOI: 10.3897/asp.81.e98166

Authors: Xiao-Zhu Luo, Mariam Gabelaia, Arnaud Faille, Rolf Beutel, Ignacio Ribera, Benjamin Wipfler

Abstract: The stable environment of subterranean realms is characterized by constant darkness, temperature and humidity, and scarcity of resources. This led to similar adaptations in different lineages of animals, such as the reduction of eyes and pigmentation. It is common textbook knowledge that blindness in cave insects is compensated for by transformations of other sensorial structures, especially the antennae with their rich array of sensilla. We tested this hypothesis with 33 species of Leiodidae of the tribe Leptodirini (Coleoptera) with and without eyes and from hypogean and epigean environments. We documented and compared the number, types, arrangement and density of smooth and furrowed antennal sensilla on certain flagellomeres. Our statistical analysis that took effects of body size and phylogeny into consideration showed that (1) the number of these sensilla does not differ between hypogean or epigean beetles; (2) the same applies to length and diameter of the antennal sensilla; (3) there is a difference in density, but unexpectedly it is lower in hypogean species. Our finding thus contrasts with widely accepted earlier interpretations for those external antennal sensilla in the studied Leptodirini, showing that sensillar patterns are scarcely affected in these subterranean beetles if at all, and even less dense in blind and cave-living species. Our results thus add a new facet to the evolution of cave animals.

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DOI: 10.3897/asp.81.e107507

Authors: Elton Orlandin, Mônica Piovesan, Daniel Herbin, Eduardo Carneiro

Abstract: Apatelodidae is a family of Neotropical bombycoids that still needs to be studied in several aspects, as many groups of species have not yet undergone a careful systematic revision. On the other hand, recent studies showed that some species known to be widely distributed in fact form species complexes. Until now, Olceclostera bifenestrata Schaus, 1912, described from Costa Rica, supposedly has a wide distribution, reaching the south of Brazil. We reviewed specimens from South America identified as O. bifenestrata. Using morphological data and DNA barcodes, we discovered that South American specimens belong to four new species: Olceclostera jairana sp. nov., Olceclostera quilombola sp. nov., Olceclostera xeta sp. nov. and Olceclostera wayana sp. nov. The four species can be distinguished mainly by characters of genitalia, mostly in aedeagus structures. However, wing characters (number of hyaline spots) and abdomen characters (number of patches of larger scales) may be useful for classifying species groups in Olceclostera. Additionally, we illustrate and describe for the first time all the life stages of an Olceclostera species, including its chaetotaxy and life history, and provide the first SEM images of the egg of an Apatelodidae species. We present a discussion about the morphological characters of adults and immatures and their relevance to the systematics of Apatelodidae.

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DOI: 10.3897/asp.81.e106356

Authors: He-Nan Li, Wen-Ya Pei, Ming-Fu Wang, Bang-Qing Chen, Hong-Lin Peng, Rong-Jun Cao, Ming-Teng Zhao, Jun Yang, Xiao-Chen Zhang, Dong Zhang

Abstract: Anthomyiidae is a cosmopolitan and diverse family of Calyptratae, and is routinely considered to play key roles in both ecology and agriculture. The higher-level phylogenetic classification of Anthomyiidae has been highly controversial, necessitating further molecular data for precise reconstruction of phylogenetic relationships. In this study, we successfully acquired and annotated 18 new mitogenomes of anthomyiids. Moreover, the mitogenomes of the following genera Eustalomyia, Hyporites, Leucophora, Emmesomyia and Eutrichota are reported for the first time. The 18 mitogenomes are compared with confamilial species to assess genetic variation and to better understand evolutionary relationships within the family Anthomyiidae. In comparisons among 13 mitochondrial protein coding genes (PCG), the calculation of evolutionary rate exhibited nad1 as the fastest evolving gene in Anthomyiidae. Among the anthomyiids investigated, cox2 and nad4 had the lowest genetic distance across the 13 PCGs, suggesting a high degree of conservation for these two genes. Herein, we conducted phylogenetic analyses of the newly sequenced mitogenomes along with 11 known anthomyiids to investigate the interrelationships of Anthomyiidae. Our results indicate that Anthomyiidae is a monophyletic lineage and sister group to Scathophagidae, confirming prior findings based on morphological and molecular analyses. We recovered two subfamilies as monophyletic (Myopininae, Pegomyinae) while Anthomyiinae was polyphyletic. The great species diversity of anthomyiid flies limits the availability of mitogenomes for accurately resolving the phylogeny of Anthomyiidae. Nonetheless, our study provides novel insight into the molecular taxonomy, evolution, and phylogeny of the family Anthomyiidae.

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DOI: 10.3897/asp.81.e102928

Authors: Andrea Basso, Dimitrios Avtzis, Christian Burban, Carole Kerdelhué, Kahraman İpekdal, Emmanuelle Magnoux, Jérôme Rousselet, Enrico Negrisolo, Andrea Battisti

Abstract: The pine processionary moth, Thaumetopoea pityocampa, is an iconic insect in the Mediterranean culture because of its economic and medical importance and the unique traits of the life history, namely the winter feeding and the construction of conspicuous silk tents by the larvae. Its taxonomic status, however, is unclear because the type material is not available and there is confusion among the several species and subspecies described in the last centuries. In the present study, a metadata analysis of morphological and molecular data of the species in the T. pityocampa complex were considered for species delimitation, using more than 400 individuals from more than 120 geographic locations. A reconstruction of the origin of the material used for the first formal description has allowed to identify the type locality and to design a neotype. In addition, as Denis and Schiffermüller were referring to the work of Réaumur for details about the species, the description provided by Réaumur was reviewed. The results indicate that the barcode region of mitochondrial DNA is a reliable trait to separate species in most cases whereas morphological traits are not. Hybridization among taxa makes it difficult to delimit species in contact zones when mating barriers are not present. In other cases, such as the populations of Crete Island, the lack of gene flow with the mainland population may support species delimitation even when morphological traits are not conclusive. Thus, the new species Thaumetopoea cretensis is described here based on the evidence obtained from a previous study. Species delimitation based on both mitochondrial and nuclear markers allowed maintenance of three species of the complex (T. cretensis, T. pityocampa, and T. wilkinsoni) while more data are needed to determine the status of two recently described species: T. hellenica and T. mediterranea.

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DOI: 10.3897/asp.81.e101438

Authors: Thomas J. Simonsen, Marie Djernæs, Ole Fogh Nielsen, Kent Olsen

Abstract: We explore haplotype diversity, phylogeography and phylogenetic relationships of the damselfly Platycnemis pennipes in Europe based on 618 bp DNA from the mitochondrial gene COI. A haplotype network analysis shows that the species is divided into two haplotype groups. One is restricted to the Italian Peninsula, while the other is found from the Black Sea region across eastern and central Europe to Scandinavia, England, and southwestern France. This pattern is recovered in a Bayesian phylogenetic analysis. Genetic distance (K2P) between the two groups is approximately 1.5%, while within-group variation is an order of magnitude lower. An analysis of the molecular variance (AMOVA) shows that variation between the two groups account for more than 96% of the total variation within the dataset, adding to the evidence that they have been isolated for a considerable amount of time. The pattern we find is similar to the so-called Grasshopper Paradigm in European phylogeography, where a species has recolonized Europe after the last glaciation from a glacial refugium in the southeast, while other refugial populations in the Iberian and Italian peninsulas have remained isolated to this day. In P. pennipes there is only an isolated refugial population in Italy as the species does not have current populations in the Iberian Peninsula. By comparing the genetic distance between the two groups to a previously published divergence time analysis of European Odonata we estimate that they have likely been isolated since the onset of the Saale Glaciation ca 400 ky ago.

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DOI: 10.3897/asp.81.e104810

Authors: Zachary H. Griebenow, Adrian Richter, Thomas van de Kamp, Evan P. Economo, Ziv E. Lieberman

Abstract: The male genitalia of the Insecta are famed for structural and functional diversity. Variation in this anatomical region shows ample phylogenetic signal, and this variation has proven indispensable for classification across the insects at multiple taxonomic ranks. However, in the ants (Hymenoptera: Formicidae) the male genital phenotype is ancillary to the morphology of the worker caste for systematic purposes. Ants of the enigmatic subfamily Leptanillinae are an exception, as males are easier to collect than workers. Ongoing systematic revision of the Leptanillinae must therefore rely upon the male phenotype – particularly the spectacular morphological profusion of the male genitalia. To thoroughly illuminate this anatomical region and aid comparative morphological research on ant male genitalia, we present a comparative morphological study of the male genitalia in nine exemplar lineages spanning the Leptanillinae, plus three outgroups representing other major clades of the Formicidae. We use micro-computed tomography (micro-CT) to generate 3D volumetric reconstructions of male genital skeletomusculature in these specimens. Our descriptions use new muscular terminology compatible with topographic main-group systems for the rest of the pterygote soma, and applicable to all Hymenoptera. We find that male genitalia in the Leptanillinae show an overall trend towards skeletomuscular simplification, with muscular reduction in some cases being unprecedented in ants, or even hymenopterans in general. In several lineages of the Leptanillinae we describe derivations of the male genitalia that are bizarre and unparalleled among the Hymenoptera. We conclude by discussing the functional implications of the often-extreme morphologies here observed.

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DOI: 10.3897/asp.81.e104426

Authors: Noor Izwan-Anas, Van Lun Low, Zubaidah Ya’cob, Emmanuel Y. Lourdes, Mohamad Rasul Abdullah Halim, Mohd Sofian-Azirun, Hiroyuki Takaoka, Peter H. Adler

Abstract: Black flies play a prominent role in public health and the epidemiology of parasitic diseases of humans, domesticated and wild animals. Correct identification and a comprehensive survey are required to identify vector and pest species and thus understand their biological attributes which play a vital role in the monitoring program. DNA barcoding is an established molecular tool that provides rapid and accurate species identification. Our study strengthens the molecular database for black flies in Malaysia by adding 59 cytochrome c oxidase I sequences for 22 species, of which 14 are included for the first time. These sequences, combined with those in public databases, represent a total of 338 sequences for 52 Malaysian species, nearly 50% of which were collected from type localities. At the subgeneric level, barcode gap analysis most accurately identified species in the subgenus Nevermannia (92%), followed by Simulium s. l. (91%), and Gomphostilbia (81%). The remaining sequences were ambiguous and could not be distinguished from those of nearest neighbour species due to an overlap in genetic divergence and low genetic diversity, especially between insular species. Tree analyses indicate that certain species had incomplete lineage sorting and low mitochondrial signals. Possible cryptic species were indicated in the Simulium (Gomphostilbia) batoense and S. (G.) epistum species groups. Species delimitations were consistent with morphological identifications except in large species groups such as the S. (G.) asakoae, S. (G.) batoense, S. (G.) epistum, and S. (Simulium) melanopus groups. The use of type specimens or specimens collected from type localities (topotypes) in barcoding is strongly recommended for reference sequences to increase the reliability of the molecular database.

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DOI: 10.3897/asp.81.e110014

Authors: Steffen U. Pauls, Wolfram Graf, Anna E. Hjalmarsson, Alan Lemmon, Emily Moriarty Lemmon, Malte Petersen, Simon Vitecek, Paul B. Frandsen

Abstract: Streams represent a special case of directional environmental gradients where ecological opportunity for diversification may be associated with upstream and downstream dispersal into habitats that differ in selective pressures. Temperature, current velocity and variability, sediment erosion dynamics and oxygen saturation are key environmental parameters that change in predictable ways from springs to river mouth. Many aquatic insects occupy specific longitudinal regions along these gradients, indicating a high degree of adaptation to these specific environmental conditions. In caddisflies, the evolution of tracheal gills in larval and pupal stages may be a major driver in oxygen uptake efficiency and ecological diversification. Here we study the evolution of larval gill structure in the Rhyacophila vulgaris species group using phylogenomic methods. Based on anchored hybrid enrichment, we sequenced 97 kbp of data representing 159 independent nuclear protein coding gene regions to infer the phylogeny of the R. vulgaris species group, whose species exhibit both high diversity of gill types and varied longitudinal preferences. We find that the different gill types evolved independently as derived characters in the genus and that gill structure is linked to the longitudinal habitat preference, thereby serving as a possible ecological key innovation in the R. vulgaris group.

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DOI: 10.3897/asp.81.e111281

Authors: Hao Yu Liu, Ruo Lan Du, Wei Zhao, Xing Ke Yang, Yu Xia Yang

Abstract: Insect male genitalia show an evolutionarily variable morphology that is valuable for both species identification and phylogenetic analyses. However, we often encounter some difficulties when conducting relevant studies due to only quantitative variations exhibited in male genitalia. In this study, based on the taxonomy of the Macrolycus dotatus species group (a total of seven species, including three new species described here), we analyzed the male genitalia shapes by GM and then constructed the phenotypic relationships by UPGMA, NJ and MP analyses. The results demonstrated that the species could be well delineated by the shape of male genitalia, and the produced phenograms frequently recovered phenotypic similarity between the coupled species, including M. atronotatimimus sp. nov. + M. huoditangensis sp. nov. and M. aemulus + M. dotatus, which is useful for making comparisons in species descriptions. Meanwhile, the MP analysis of male genitalia shape using two landmark configurations is considered reliable in inferring the phylogenetic relationship among species in terms of the consistency between its topologies and the molecular phylogeny. This study sheds new light on improving the morphological taxonomy of insects in lower grades while fully utilizing the taxonomic value of male genitalia in a phylogenetic context.

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DOI: 10.3897/asp.81.e96839

Authors: Sana Sharifian, Ehsan Kamrani, Maria A. Nilsson, Hanieh Saeedi

Abstract: Brachyuran crabs constitute the dominant fauna in intertidal and supratidal coasts of mangrove forests. We sampled the most commonly occurring crab species from the biodiversity rich Persian Gulf mangrove forest. We identified crabs from Camptandriidae, Dotillidae, Ocypodidae, Macrophthalmidae, and Sesarmidae as the most common species in the sampled regions. Molecular barcoding was applied to determine 11 species (29 specimens) of the mangrove crabs. Two mitochondrial genes were used to barcode the specimens and these were included in a larger phylogenetic data set. Many of the analyzed species showed a close phylogenetic relationship with species from the Northern Arabian Sea. The results provide the first steps to study the genetic diversity of the mangrove crab community along the Iranian coasts to support protection and management of mangrove ecosystems and its associated taxa.

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DOI: 10.3897/asp.81.e109833

Authors: Pavel Sroka, Jakub Prokop

Abstract: The stonefly suborders Arctoperlaria and Antarctoperlaria reflect the current division of the diversity of this insect order between the Northern and Southern Hemispheres. However, there are several exceptions to this pattern, the most notable being the family Notonemouridae, which is phylogenetically deeply subordinate within the northern Arctoperlaria, but distributed in South Africa, South America, and Australia. Various hypotheses have been proposed regarding the circumstances of their dispersal to the south. Some estimated their origin as relatively recent, with long-distance dispersal to the southern continents in the Late Cretaceous or early Paleogene. On the other hand, fossils of Notonemouridae have been dated to the Middle Jurassic, proving the lineage is very ancient. However, all known notonemourid fossils originate from Asia and the timing of their dispersal to the south cannot be precisely estimated. Here we report new fossil stoneflies from the Late Jurassic Talbragar Fish Beds, Australia, described as Talbragaria australis gen. et sp. nov. and attributed to Notonemouridae. This finding represents the first fossil evidence of the northern suborder Arctoperlaria in the Southern Hemisphere, and confirms the north-to-south dispersal of Notonemouridae across Pangea prior to the continental break-up.

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DOI: 10.3897/asp.81.e104396

Authors: Fedor V. Konstantinov

Abstract: Plant bugs of the genus Heterocapillus (Miridae: Phylinae) are predominantly Mediterranean minute black phylines with distinctly swollen antennomeres I and II. The monophyly and relationships of this group has been questioned in several studies. This paper provides a morphology-based phylogeny inferred with Parsimony and Bayesian analyses of the genus and a diverse array of outgroups. Based on these results, Heterocapillus is here redefined as monotypic. A new monotypic genus Abietocapsus gen. nov. is erected to accommodate Phoenicocoris dissimilis (Reuter, 1878) and updated diagnoses are provided for the genera Lobicris Putshkov, 1977 gen. dist., Mesopsallus Wagner, 1970 gen. dist., and Salicarus Kerzhner, 1962. The following new combinations are established: Campylomma atlanticum (Wagner, 1963) comb. nov., Campylomma pusillum (Reuter, 1878) comb. nov., Mesopsallus amygdali (Wagner, 1960) comb. nov., Mesopsallus fagi (Drapolyuk, 1990) comb. nov., Mesopsallus mali (Meyer-Dür, 1843) comb. nov., Mesopsallus pici (Reuter, 1899) comb. nov., Mesopsallus rhodani (Fieber, 1861) comb. nov., Mesopsallus validus (Reuter, 1901) comb. nov., Psallus (Phylidea) validicornis (Reuter, 1876) comb. nov., Salicarus cavinotum (Wagner, 1973) comb. nov., Salicarus genistae (Lindberg, 1948) comb. nov., Salicarus nitidus (Horváth, 1905) comb. nov., and Salicarus perpusillus (Wagner, 1960) comb. nov. Atractotomus schmiedeknechti Reuter, 1899 is synonymized with Atractotomus parvulus Reuter, 1878. Heterocapillus niger Wagner, 1966 is synonymized with Psallus (Phylidea) validicornis (Reuter, 1876). The highly homoplastic nature of the antennal shape in phylines is demonstrated, suggesting against using these features as diagnostic at the generic level.

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DOI: 10.3897/asp.81.e107579

Authors: Si-Xun Ge, Zhuo-Heng Jiang, Li-Li Ren, Cornelis van Achterberg, Jiang-Li Tan

Abstract: The family Stephanidae (Hymenoptera) constitutes a unique group within the Apocrita, playing a pivotal role in the evolution of parasitoid wasps. Although the phylogeny of Stephanidae has been previously inferred, it remains at a low resolution when considering both extinct and extant genera, as well as the enigmatic extinct genus †Electrostephanus. Here, we undertake a revision of Stephanidae extinct, presenting descriptions of new specimens from late Cretaceous Burmese amber and early Eocene Baltic amber. Combining all extant and extinct genera, the phylogeny of Stephanidae was analyzed, incorporating 57 species within 21 genera based on 64 morphological characters. We apply both under maximum parsimony with equal weighting and implied weighting methods, with four species representing early Apocrita as outgroups. Divergence times are estimated by utilizing extinct taxa as calibration points. A new basal subfamily of stephanid wasp, †Lagenostephaninae subf. nov. was established, encompassing †Lagenostephanus and the newly described genera †Tumidistephanus gen. nov and †Neurastephanus gen. nov. The genus †Electrostephanus is redefined, with two species assigned under distinct genera, †Neurastephanus gen. nov. and †Aphanostephanus gen. nov.. We discuss some of the putative morphological synapomorphies of evolutionary significance within the phylogenetic framework. Our results complement several characteristics of great taxonomic importance for Stephanidae and provide new insights into the early evolution of the family.

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DOI: 10.3897/asp.81.e106948

Authors: Marco Isaia, Giuseppe Nicolosi, Alessandro Infuso, Carles Ribera

Abstract: The spider genus Typhlonesticus Kulczyński, 1914 (Araneae: Nesticidae) includes seven species, most of which exhibiting strict preference for caves, abandoned mines and other subterranean habitats. In Italy the genus is represented by two species: T. morisii (Brignoli, 1975), an extremely narrow endemic species from SW-Alps with a very high level of subterranean adaptation and T. idriacus (Roewer, 1931), showing a much wider distribution in NE-Italy and poor adaptations to subterranean life. Our recent biospeleological surveys in the Alps lead to the discovery of new populations of highly troglomorphic Typhlonesticus that proved to belong to two new species based on morphological and molecular data. Considering the rarity of these new species, we provide general information on their ecology and distribution, including a comparative analysis of troglomorphic traits in Typhlonesticus in relation to biogeographic factors. Information on the conservation status, useful for assessing their extinction risk based on International Union for Conservation of Nature (IUCN) guidelines, is also provided.

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DOI: 10.3897/asp.81.e104522

Authors: Clément Schneider, Cyrille A. D’Haese

Abstract: Mackenziella psocoides Hammer, 1953 (Collembola: Mackenziellidae) is a widespread but uncommon springtail. Its unusual body shape (ovoid, with partial coalescence of abdominal segments) has puzzled the specialists for a long time, until the discovery of males allowed to relate the species to a family of globular springtails, the Sminthurididae. Yet, the precise phylogenetic position of M. psocoides, and hence of the Mackenziellidae, remained ambiguous. In this work, we report a new locality for M. psocoides in Germany. We provide the first DNA sequences (nuclear ribosomal DNA operon) for the species, as well as the first images using scanning electron microscopy. We investigate its phylogenetic position based on the molecular data and specify details on its morphology. Our results show that M. psocoides is nested inside of Sminthurididae, as the sister group of Sphaeridia Linnaniemi, 1912. Consequently, Mackenziellidae syn. nov. is here synonymized with Sminthurididae. We include Mackenziella and Sphaeridia in the Sphaeridiainae subfam. nov., a replacement name for Sphaeridiinae Richard, 1968 that is a junior homonym of Sphaeridiinae Latreille, 1802 (Coleoptera: Hydrophilidae). Corresponding to its phylogenetic position within Sminthurididae, the evolutionary origin of M. psocoides is younger than previously thought (79 mya +/- 35 my). The lineage accumulated an unusual amount of body modifications involving, among others, the loss of the globular body shape. This rapid rate of evolution is, to our knowledge, unique in springtails. It shows that globular body shape is not an evolutionary dead-end, and the secondary acquisition of a linear body shape and recovery of longitudinal flexibility is still possible.

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DOI: 10.3897/asp.81.e104772

Authors: Yan-Na Zheng, Jun-Jie Gu, Zhu-Qing He, Huateng Huang, Li-Bin Ma

Abstract: The hemispherical head is prevalent in Gryllinae crickets, so the rare crickets that have a unique form of head will be extremely unusual. In previous studies, this special feature can be one of the important features to distinguish and identify these crickets. But does this particular head shape truly reflect a clear-cut taxonomic relationship? The species of the genus Loxoblemmus have a typical truncate head; species of the genus Stephoblemmus have a more exaggerated truncate head, with the frontal end even extending into a lamellar. The genus Mitius is relatively unusual in that species of this genus have both globose or truncate heads. How are these species related? Does the cephalic shape perfectly reflect the natural classification of these species? Based on these questions, the study applied species definition and morphological classification to explore the intergeneric and intrageneric species relationships of the genera Mitius, Stephoblemmus, and Loxoblemmus, and derived the following main conclusions: (1) Mitius and Stephoblemmus are related and distinct from Loxoblemmus; (2) Mitius species bear two types of frons (truncated and rounded), but this feature disallows them to be classified as natural groups; (3) one genus synonym and three species synonyms are raised (Mitius Gorochov, 1985 syn. n., Mitius splendens (Shiraki, 1930) syn. n., Mitius eryuanensis Yuan, Xie & Liu, 2021 syn. n. and Mitius brevipennis Yuan, Ma & Gu, 2022 syn. n.), and seven new status combinations are proposed (Stephoblemmus blennus (Saussure, 1877) comb. n., Stephoblemmus castaneus (Chopard, 1937) comb. n., Stephoblemmus enatus Gorochov, 1994 comb. n., Stephoblemmus flavipes (Chopard, 1928) comb. n., Stephoblemmus minor (Shiraki, 1911) comb. n., Stephoblemmus minutulus (Yang & Yang, 1995) comb. n. and Stephoblemmus vaturu (Otte & Cowper, 2007) comb. n.). The studies indicated that frons shapes that appear to be significantly different might not always reflect the correct Gryllinae species relationships and a combination of more taxonomic features and taxonomic techniques is needed often to reveal the true taxonomic relationships.

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DOI: 10.3897/asp.81.e105025

Authors: Hoi-Sen Yong, Sze-Looi Song, Kah-Ooi Chua, Yvonne Jing Mei Liew, I. Wayan Suana, Phaik-Eem Lim, Kok-Gan Chan, Praphathip Eamsobhana

Abstract: The complete mitogenomes of fruit flies Zeugodacus (Javadacus) calumniatus, Z. (Javadacus) heinrichi and Z. (Sinodacus) hochii have similar gene order and contain 37 genes and a non-coding region. They share an identical start codon for the respective protein-coding genes (PCGs), an identical TAA stop codon for 11 PCGs, TAG for cob, and an incomplete T stop codon for nad5. The cloverleaf structure of most of the tRNAs is similar in the three Zeugodacus species. Phylogenetic analyses reveal Z. (Parasinodacus) cilifer to be external to two main clades: (A) monophyletic subgenus Zeugodacus; and (B) subgenera Javadacus and Sinodacus. The present results indicate that the taxonomic status of some taxa needs clarification. Z. calumniatus is genetically very similar to Z. tau and is not congruent with its current placement in the munda complex. Z. mukiae NC_067083 is genetically very similar to Z. scutellaris, but differs significantly from Z. mukiae MG683384 of the arisanicus (arisanica) complex. On the other hand, Z. proprediaphorus is genetically distinct from and not a synonym of Z. diaphorus. Z. caudatus sensu stricto from Indonesia forms a sister lineage with Z. diversus, instead of with the Malaysian and Chinese taxa of Z. caudatus sensu lato. A notable incongruence is the sister lineage of Z. (Sinodacus) hochii and Z. (Javadacus) heinrichi among other taxa of subgenus Javadacus. A more extensive taxon sampling, particularly the subgenus Sinodacus (and other subgenera), is needed to clarify/resolve their subgenus status.

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DOI: 10.3897/asp.81.e97946

Authors: Zehui Kang, Yuanyuan Xu, Guoquan Wang, Ding Yang, Xiao Zhang

Abstract: Limoniidae, the most speciose family in the superfamily Tipuloidea, consists of four subfamilies and more than 11,000 species. However, mitochondrial (mt) genome sequences, which have been widely used for phylogenetic study, are available for only 11 species across three subfamilies. Thus, a larger variety of mt genome sequences in Limoniidae are required to improve our understanding of tipuloid phylogeny and genomic evolution. Here we present mt genomes of Elephantomyia (Elephantomyia) inulta Alexander, 1938 and Helius (Helius) pluto Alexander, 1932, representing the first mt genomes of the tribe Elephantomyiini (Limoniidae). The two mt genomes are typical circular DNA molecules and show similar gene order, nucleotide composition and codon usage. Standard ATN start and TAR stop codons are present in most protein-coding genes. All transfer RNA (tRNA) genes exhibited the cloverleaf secondary structure typical for metazoans except in tRNASer(AGN), which lacks the dihydrouridine arm. Phylogenetic analyses were performed based on four nucleotide matrixes for the currently sequenced species of Tipuloidea using Bayesian inference and maximum likelihood methods. Four-cluster likelihood mapping was used to study incongruent signals between different topologies. Pediciidae is supported as the earliest lineage in Tipuloidea, and the sister-group relationship between Cylindrotomidae and Tipulidae is also supported, but the monophyly of Limoniidae is not supported. Our study also supports the monophyly of Elephantomyiini (Elephantomyia + Helius), as one of origins of flower-visiting in Limoniidae. Although Elephantomyiini is sister to Limoniinae + Epiphragma (Limnophilinae) in our study, a more precise understanding of its phylogenetic position in Tipuloidea will require additional studies that include a broader species sample.

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DOI: 10.3897/asp.81.e98141

Authors: Ionela-Mădălina Viciriuc, Mircea-Dan Mitroiu, Richard Robinson Askew, Nicolas Ris, Lucian Fusu, Nicolas Borowiec

Abstract: The introduction of the biological control agent Torymus sinensis Kamijo (Hymenoptera, Chalcidoidea, Torymidae) to control the populations of the chestnut gall wasp Dryocosmus kuriphilus Yasumatsu (Hymenoptera, Cynipidae) is considered one of the successful programs in biological control. The species was involved in interspecific hybridisation in Japan and the specimens imported into Europe were derived from this hybrid lineage, showing signs of introgression. The discovery of mitochondrial haplotypes or possible Enolase haplotypes from T. beneficus or of specimens with shorter ovipositor does not necessarily imply that T. beneficus is present in Europe, only that the European specimens are of hybrid origin. Of the native European Torymus species associated with D. kuriphilus, the molecular and morphometric results indicate Torymus notatus (Walker) as the closest species to T. sinensis. The two are part of the same species-group (cyaneus group), are nested together in the multivariate ratio analysis and are the closest genetically based on all three nuclear markers: Enolase (1.5% divergence), Wingless (2%) and ITS2 (13%). However, on the mitochondrial marker COI the closest species is Torymus rubi (Schrank) at 9.9% divergence. As such, T. notatus is the most likely candidate for accidental interspecific hybridisation if this is to happen in Europe. We provide an illustrated identification key for the European species of Torymus associated with D. kuriphilus, an important but lacking tool for biological control programs.

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DOI: 10.3897/asp.81.e101374

Authors: Kateřina Rosová, Jakub Prokop, Jörg U. Hammel, Rolf G. Beutel

Abstract: Omophroninae is a distinctive monogeneric group of Carabidae, presumably placed relatively close to the root of the megadiverse adephagan family. In the present study we describe a larva belonging to Omophroninae from mid-Cretaceous Burmese amber and erect a new genus †Cretomophron. Several features support the placement in this small but distinctive subfamily, such as the wedge-shaped head, the large triangular nasale, the elevated antennae with the apical segment directed sideways, the large and bidentate mandibular retinaculum, the enlarged hexagonal prothorax, legs with a distinct armature of spines, and the relatively narrow and posteriorly tapering abdomen. In contrast to larvae of the extant genus Omophron Latreille, the posterior tentorial grooves are not shifted backwards, apparently a plesiomorphic feature, the 2nd antennomeres are markedly longer, and the legs bear long setae and rather thin and long spike-like setae. †Cretomophron also differs in the presence of numerous setae arranged in transverse rows on abdominal segment VI. Lateral lobe-like expansions of abdominal tergites are a conspicuous feature of the new genus but similar structures occur in later instars of Omophron. Structural specializations of the head, prothorax and legs strongly suggest that the larvae were burrowing in sand, like adults and larvae of the extant genus, and that they were efficient predators, detecting prey with the unusually shaped antennae and long maxillae, grasping it with the elongate apical mandibular tooth, and squeezing and piercing it between the bidentate retinaculum and large and triangular nasale.

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DOI: 10.3897/asp.81.e102848

Authors: Nathalia H. Pecly, Daniela M. Takiya, Rodney R. Cavichioli, Gabriel Mejdalani

Abstract: The genus Dasmeusa is distributed in Northern and Northeastern Brazil and the Guianas. Until the present study, six species were recognized within Dasmeusa. The species of this genus are very similar in terms of color and external morphology, being distinguished mainly by the male terminalia. Here, we review and redescribe Dasmeusa and its species, describe four new species, and present the first phylogenetic analysis of the genus, including 40 morphological characters and 15 terminal taxa. Dasmeusa flavescens Metcalf and Erythrogonia bicolor Metcalf are considered junior synonyms of the type-species, Dasmeusa pauperata (Fabricius). Scanning electron microscopy was employed for a detailed study of the integument of the type-species, including sensilla, surface sculpturing, brochosomes, organ of Evans, and other structures. The phylogenetic analysis with equal weights resulted in nine most parsimonious trees. The implied weighting method resulted in two trees, both with the same ingroup topology as observed in one of the nine equal-weights trees. This preferred topology is as follows: ((D. basseti (D. mendica (D. rafaeli sp. nov., D. falcifera sp. nov.))) (D. isabellina (D. oriximina sp. nov. (D. pauperata (D. imperialis, D. dinizi sp. nov.))))). Dasmeusa was recovered as monophyletic in all trees, being supported by five apomorphic characters.

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