DOI: 10.3897/asp.84.e197461

Authors: Alicja Pełczyńska, Evgeny E. Perkovsky, Brendon E. Boudinot, Adrian Richter, Wiesław Krzemiński, Tomasz Kołodziej, Pawel Korecki, Katarzyna M. Sowa, Agnieszka Soszyńska

Abstract: Abstract The extinct Eocene fly genus Kelneria Matile, 1979 is characterized by extreme modification of male genitalia within the family Keroplatidae. However, their morphology has remained poorly understood, as fossil material cannot be examined using traditional entomological techniques. To overcome this limitation, we applied micro-computed tomography (µCT) and synchrotron radiation micro-computed tomography (SRµCT) to reconstruct the genitalia through segmentation of scan data, enabling reassessment of the spatial organization and homology of their structures. Our analyses indicate that Kelneria was both morphologically distinctive and well represented in Eocene amber assemblages. They also support the synonymization of the Oise amber genus Eokelneria Hebert, Ngô-Muller & Nel, 2025 with Kelneria Matile, 1979, demonstrating that the characters used to establish Eokelneria fall within the range of variation observed in Kelneria. We further demonstrate that the species-level diversity of Kelneria has been underestimated and describe three new species: Kelneria rovnensis Pełczyńska & Perkovsky sp. nov., K. erroris Pełczyńska, Krzemiński & Soszyńska sp. nov. and K. szymoni Pełczyńska, Krzemiński & Soszyńska sp. nov. Additionally, identification of representatives in Rovno and Oise ambers extends the geographic and temporal range of the genus to the early Eocene (c. 53 Ma). Taken together, these findings show that Kelneria was an abundant, diverse, and widespread genus in the Eocene but has no modern representatives. The reasons for this extinction remain unknown; however, contrasting distribution of Eocene and extant representatives of the tribe Robsonomyiini suggests that this disappearance may reflect a broader faunal turnover in Europe after the Eocene.

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DOI: 10.3897/asp.84.e189651

Authors: Monika Pramatarova, Daniel Burckhardt, Igor Malenovský, Ilia Gjonov, Albena Lapeva-Gjonova

Abstract: Abstract Cuticular microsculpture is an important morphological feature in insects, but it has not been systematically investigated in psyllids (Hemiptera: Psylloidea). Here, we examine the microsculpture of the vertex and thoracic dorsum in two subfamilies of Aphalaridae, focusing mainly on European taxa and using scanning electron microscopy to evaluate its taxonomic and phylogenetic significance. In Aphalarinae, imbricate microsculpture was observed in Aphalara, Eumetoecus and Rhodochlanis; rugose microsculpture in most species of Colposcenia; and several microsculptural types in Craspedolepta, including imbricate, smooth–imbricate, areolate–rugose, and alveolate patterns. A unique tuberculate microsculpture was found in Rhinocola aceris (Linnaeus, 1758), whereas other Rhinocolinae exhibited an imbricate pattern. Ancestral state reconstruction indicates that imbricate microsculpture represents the plesiomorphic condition in Aphalaridae and remains expressed to varying degrees in most extant representatives of the family. Multiple transitions among sculptural types within Craspedolepta and partly also Colposcenia indicate that microsculpture can evolve relatively rapidly, although the distribution of character states across the phylogeny shows that it still retains phylogenetic signal at shallow to intermediate taxonomic levels, despite a moderate level of homoplasy. Overall, these results demonstrate that head and dorsal thoracic microsculpture constitute diagnostic and phylogenetically informative characters that can be more widely applied in taxonomy, systematics and evolutionary research on psyllids.

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DOI: 10.3897/asp.84.e174408

Authors: Shao-Li Mao, Hao Yuan, Xuan-Zeng Liu, Yan-Wen Wang, Lu-Yao Yang, Ya-Fu Zhou

Abstract: Abstract The tribe Meconematini (Meconematinae) is a species-rich lineage whose genus- and species-level classifications remain taxonomically controversial. In this study, we obtained 17 new mitochondrial genomes using high-throughput sequencing and conducted the first comparative genomic and phylogenomic analyses within this tribe. The mitogenomes of Meconematini exhibit conserved structural features typical of Meconematinae, including conserved gene order, pronounced AT bias, and negative GC-skew. Notably, a rare mitochondrial initiation codon (GTG) was identified in the ATP6 gene in Meconematini. Evolutionary analyses indicated that strong purifying selection has dominated the evolution of Meconematini mitogenomes. Phylogenetic reconstructions strongly supported the monophyly of the genera Microconema Liu and Decma Gorochov, as well as the subgenus Eoxizicus Gorochov. In contrast, the genera Phlugiolopsis Zeuner, Xizicus Gorochov, and Xiphidiopsis Redtenbacher were not recovered as monophyletic. Additionally, the phylogenetic results support the recognition of Eoxizicus and Euxiphidiopsis Gorochov as distinct genera. Importantly, mitogenomic divergence patterns were correlated with genitalia structural traits: genera within the proximal clade possess membranous male genitalia (hagloid type), whereas those in more basal clades exhibit partially or fully sclerotized genitalia (tettigonioid or grylloid type). These findings collectively enable the proposal of taxonomic revisions for the non-monophyletic genera and establish a mitogenomic baseline for reconciling morphological convergence with evolutionary relationships in Meconematini.

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DOI: 10.3897/asp.84.e174388

Authors: David Honsberger, Gabriela Pérez-Lachaud, Karl N. Magnacca, Jacob Hauʻoli Lorenzo-Elarco, Sandor L. Kelly, Mark G. Wright, Tara D. Gariepy

Abstract: Abstract Despite the recorded presence in Hawai'i of Cephalonomia hyalinipennis, a presumed parasitoid of the coffee berry borer (Hypothenemus hampei), it has never been recovered from infested coffee berries. Here we show, using morphometric and molecular data, that what has been recorded in Hawai'i as C. hyalinipennis is actually a complex of morphologically similar species, each distinct from specimens attacking the coffee berry borer in Chiapas, and from the C. hyalinipennis types from Florida. The taxon from Chiapas utilizing the coffee berry borer is also found to be distinct from C. hyalinipennis sensu stricto, and is here described as C. soconuscensis sp. nov. The five species found in Hawai'i are described as Cephalonomia lynnea sp. nov., a parasitoid of Hypothenemus eruditus; Cephalonomia luiji sp. nov.; Cephalonomia liula sp. nov., a parasitoid of a Leptophloeus sp. known to predate on the coffee berry borer; Cephalonomia kala sp. nov.; and Cephalonomia zander sp. nov., a parasitoid of an Ozognathus sp. Laboratory testing showed that C. lynnea has the ability to develop on coffee berry borer larvae, but it does not appear to do so in the wild. In addition to morphological data, DNA barcoding was used to provide further support for distinguishing species in the C. hyalinipennis species complex. The large dataset of morphometric measurements showing reliability of different ratios, and degenerate PCR primers developed to permit assembly of COI barcodes even from old museum specimens, should enable more rapid discernment within such cryptic species complexes in the future.

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DOI: 10.3897/asp.84.e168722

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

Abstract: Abstract Three Sicilian populations belonging to the enigmatic genus Xerobiotus were analyzed using integrative taxonomic methods combining detailed morphological and genetic data. A new population of X. inermis was found at its locus typicus in the coastal dunes of Gela, Sicily, while X. euxinus was recorded for the first time in Sicily at Viagrande (Catania) and Serra La Nave (Etna). Both species were examined using phase contrast microscopy (PCM) and scanning electron microscopy (SEM), and sequenced for four molecular markers: 18S rDNA, 28S rDNA, ITS-2, and COI. For X. inermis, the complete mitochondrial genome was also obtained. These data enabled us to redescribe X. inermis and to re-evaluate and amend the description of the widespread and morphologically variable X. euxinus. To explore phenotypic and quantitative variation, we conducted a principal component analysis (PCA) of morphometric traits across Xerobiotus populations and selected Macrobiotus taxa. The analysis revealed extensive character overlap among species, limiting diagnostic resolution and complicating genus-level delimitation. Phylogenetic reconstructions and species delimitation analyses were partially congruent with earlier studies but also challenged the current boundaries of Xerobiotus, especially with Macrobiotus, suggesting potential synonymy or misclassification among genus- and species-level taxa. In this paper, we integrate multiple lines of evidence from our analyses and comparisons in order to synonymize two genera and two species and identify two additional taxa as species in need of urgent revision to test for further potential synonymies.

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DOI: 10.3897/asp.84.e180526

Authors: Sarah Gerken, Magdalena Błażewicz, Kevin M. Kocot, Stefan Richter, Martin Schwentner, Regina Wetzer

Abstract: Abstract Peracarida (Arthropoda: Crustacea) is one of Earth’s most strikingly diverse animal groups. Often compared with their relatives, the decapods (crabs, shrimp, and lobsters), a better comparison would be with the hyperdiverse insects: small bodies, exceptional diversity of form, and pervasive habitat occupancy. Those traits, and that most live in the challenging ocean environment, have left the peracarids woefully underexplored. Each of the world’s few peracarid taxonomists has an extensive backlog of new species waiting to be described, and every field trip that targets peracarids yields novel species – the scope for diversity discovery is nearly unlimited. The historically sparse diversity sampling has left our understanding of peracarid relationships in a murky state. Making sense of the group’s diversity requires a solid phylogenetic framework to explain their evolutionary history. Here we summarize the 180 years of peracarid history with references to all key taxonomic discoveries and hypotheses. Beyond bringing a historical perspective to the group, we propose phylogenomic approaches to deciphering peracarid phylogeny enabled by current international projects. We welcome collaboration with all researchers working on Peracarida and are particularly interested in partnerships that broaden taxon sampling, expand geographic and habitat coverage, and support opportunities and training for the next generation of peracarid researchers.

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DOI: 10.3897/asp.84.e181309

Authors: Zhuo Chen, Hu Li, Wanzhi Cai

Abstract: Abstract The thread-legged bug genus Chinemesa Wygodzinsky, 1966 (Hemiptera: Heteroptera: Reduviidae: Emesinae: Emesini) is a small group of five species endemic to the Oriental Region. However, the species diversity of this genus is still far from completely explored, especially in the mainland of Asia. Here we describe three new species, C. ornata sp. n., C. pulchella sp. n. and C. weilingfengi sp. n., from southern China. Based on the newly sequenced mitochondrial genomes of four Chinemesa species, we detected two gene rearrangement patterns in the genus: a translocation of trnI and trnQ in all four species, and a loss of trnW in C. pulchella sp. n. Both gene rearrangements are novel within Reduviidae as well as Heteroptera, and can be explained by the tandem duplication-random loss (TDRL) model. Phylogenetic analyses based on mitogenomic datasets recovered the monophyly of Chinemesa, with the translocation of trnI-trnQ as a potential molecular synapomorphy for the genus.

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DOI: 10.3897/asp.84.e189518

Authors: Jeremy D. Wilson, Arianna Urso, Michael G. Rix, Erich S. Volschenk, Valentina Cruz Bedón, Mark S. Harvey

Abstract: Abstract The collar-door wishbone spiders of the genus Kwonkan Main, 1983 are an Australian-endemic lineage of mygalomorph spiders that often construct elaborate burrow entrances, including collars and turrets, and remain poorly documented across their range, despite museum collections indicating high local endemism and substantial undescribed diversity. Much of the existing taxonomy, including nine of the 14 currently described species, was based on limited material and lacked modern morphological or molecular approaches to species delimitation, hindering efforts to document the remaining diversity and address conservation concerns. Here, we redescribe all nine legacy species and review Kwonkan diversity within the south-western Western Australian (SWWA) agricultural region, a highly fragmented and mostly cleared landscape harbouring extensive undescribed diversity and the threatened species K. eboracum Main, 1983. In the process, we clarify species identities, present the first molecular data for several species including K. eboracum, and describe two new species (K. elatus sp. nov. and K. yorkrakine sp. nov.) that were previously attributed to legacy species. In our review of the SWWA agricultural region fauna we identify 29 putative undescribed species and a pattern of extensive sympatry, fine-scale species turnover, and extremely restricted ranges. These findings highlight the need for continued revisionary work and potential conservation listing of additional described species such as K. wonganensis (Main, 1977).

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DOI: 10.3897/asp.84.e184638

Authors: Marcin Wiorek, David C. Lees, Niklas Wahlberg, Łukasz Przybyłowicz

Abstract: Abstract In the highly diverse, endemic Madagascan lineage of Syntomini, the “black-and-white” general body pattern appears to be present in a few closely unrelated clades (and in six existing genera), causing uncertainties in their systematics. Based on materials from five natural history collections, we confirm the monophyletic character of the morphologically variable and largely montane genus Tritonaclia Hampson, 1898, with six existing species, inferred from both molecular and morphological evidence. The taxonomy of the genus is revisited, with the description of two new species: T. vonifotsy Wiorek, sp. nov. and T. ombilahy Wiorek & Przybyłowicz, sp. nov.. Naclia melania Oberthür, 1923 syn. nov. is treated as a junior subjective synonym of Glaucopis tollini Keferstein, 1870, and lectotypes of both taxa, now placed in Tritonaclia, are designated. We provide determination keys for the genus, based on adults, and their male and female genitalia. All available data on the distribution and ecology of Tritonaclia species are presented, including new geolocalisation of some important insect collecting places in Madagascar. Additionally, we provide descriptions and illustrations of male scent scales (androconial patches) on fore- and hindwings, present in two Tritonaclia species: T. tollini and T. vonifotsy Wiorek, sp. nov. Such structures are recorded in the tribe Syntomini for the first time.

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DOI: 10.3897/asp.84.e186815

Authors: Qing Li, Gregory D. Edgecombe, Chao Jiang

Abstract: Abstract Hainanthereua albilineata gen. nov. et sp. nov. is described and illustrated based on specimens from Hainan Province, China. Morphological examination showed that these specimens belong to Thereuoneminae and share certain similarities with Thereuopodina Verhoeff, 1905. Phylogenetic reconstruction based on five genes (nuclear 18S and 28S rRNA, mitochondrial 12S and 16S rRNA, and cytochrome c oxidase subunit I) indicated that the specimens form a distinct and well-supported clade that is sister group to Thereuonema Verhoeff, 1904, so a new genus is accordingly established. Combining the evolutionary history and biogeographic framework of Thereuoneminae, this study revealed the evolutionary significance of Hainanthereua gen. nov. in the context of the Peninsular Indian Plate as a biotic ferry implicated in the origin of East and Southeast Asian lineages. Morphological similarities between Hainanthereua and Thereuopodina are symplesiomorphies of a clade that includes these two genera, Thereuonema and Thereuopoda Verhoeff, 1904.

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DOI: 10.3897/asp.84.e185344

Authors: Jędrzej Warguła, Daniel Stec, Wiktoria Dmuchowska, Michalina Krakowiak, Anastasiia Polishchuk, Magdalena Gawlak, Łukasz Kaczmarek

Abstract: Abstract African tropical forests are highly underexplored regarding microinvertebrate diversity, including tardigrades. Here, we describe two new tardigrade species from the genera Parahypsibius Gąsiorek et al., 2024 and Echiniscus C.A.S. Schultze, 1840, collected in the tropical rainforest of the Kibale National Park in southwestern Uganda. Species delimitation was based on an integrative approach combining morphological, morphometric, and molecular data. Morphological analyses employed light and scanning electron microscopy, while molecular characterization used mitochondrial COI and nuclear 18S rRNA, 28S rRNA, ITS1, and ITS2 markers. Additionally, we report Echiniscus lineatus Pilato, Fontoura, Lisi and Beasley, 2008 from Uganda for the first time, providing new sequences and reconstructing haplotype networks using previously published genetic data of this species. Phylogenetic analyses of subfamily Hypsibiinae Pilato, 1969 and genus Echiniscus clarify the relationships of the new taxa and place them within the broader phylogenetic context. Our study highlights the underestimated diversity of African tardigrades and demonstrates the importance of integrative taxonomy for species delineation. The new records from Kibale National Park provide also baseline data for understanding the distribution and phylogeny of tardigrades of the Central Africa.

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DOI: 10.3897/asp.84.e181294

Authors: Oscar M. Quispe-Colca, Nelson E. Ferretti, Juan C. Chaparro, José A. Ochoa, Rick C. West

Abstract: Abstract Recent field campaigns conducted in Peru along with the examination of museum specimens allowed us to identify small tarantulas that do not fit with any known Theraphosidae genera. Morphology and additional molecular evidence from the mitochondrial gene COI led us to propose Kiskalla gen. nov. from southern Peru, at Puno region. Three new species of Kiskalla gen. nov. (K. ignacioi sp. nov., K. yeisoni sp. nov. and K. zukuapasanka sp. nov.) are herein described, diagnosed and illustrated. Kiskalla gen. nov. differs from the known Theraphosinae genera in the presence of lateral stripes on the abdomen and a small dorsal arrowhead-shaped patch of type III urticating setae, presence of a large number of spines on all legs, short and stout setae on the dorsal metatarsi encircling the filiform trichobothria, an apical crown of long spines on the metatarsi of all legs in both sexes and scopula of tarsi IV slightly developed, restricted to the margins due to the presence of long ventral setae. Males and females show genitalic features that resemble those of Hapalotremus Simon, 1903, but differ by the development of copulatory bulb keels and the aspect of the spermathecae. We also present data on the natural history and distribution of the species.

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DOI: 10.3897/asp.84.e175149

Authors: Junta Abe, Toshiharu Mita

Abstract: Abstract Phylogenetic reconstruction of Proctotrupinae, the largest subfamily of Proctotrupidae, was conducted for the first time based on total evidence combining molecular and morphological characters. Based on our analysis, we conclude that the genus Nothoserphus, previously belonging to the tribe Cryptoserphini, should be transferred to the tribe Disogmini. New diagnostic characters for each genus were proposed based on the morphological analysis. The results of the analysis showed that the exceptionally hardened ovipositor sheath is a defining character of Proctotrupidae, and it was estimated that its morphological character states were not reflected in the phylogenetic relationship, probably because of its variability related to parasitic strategy and host habitat.

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DOI: 10.3897/asp.84.e144506

Authors: Dominic A. Evangelista, Emmy Fiorella Medina-Espinoza, Stephany Karla Mendieta Yáñez, Kim Drager, Kali L. Swichtenberg, Gillian Gile, Yifei Kang, Jiří Hromádka, Katharine Vanker, Melissa Sanchez-Herrera

Abstract: Abstract Recent systematic work has shown that Lamproblattidae may be an ancient lineage of Blattodea. Given their putative position as sister to Xylophagodea, investigations of their little-known biology can uniquely offer new insight into the evolution of wood-feeding and thus, the evolutionary precursors to eusociality. Here, we compiled all literature information on Lamproblattidae in a new appraisal, integrating it with novel data collection. First, we revised the systematics of Lamproblattidae. We provide a key to the family and related taxa, give revised definitions of taxa, remove Eurycanthablatta from Lamproblattidae, describe Lamproblatta mimadelfi sp. nov., and provide a morphology-based phylogeny for Lamproblattidae. The phylogeny weakly demonstrates the monophyly of Lamproblattidae, the paraphyly of Lamproblatta relative to Lamproglandifera, the lack of support for Eurycanthablatta within Kittrickea, and weak support for Anaplecta + Lamproblattidae. Next, we reviewed hundreds of iNaturalist observations to gain new occurrence data for Lamproblattidae. Combining this distribution data with that from a literature review and other publicly available databases, we model the habitat suitability for Lamproblattidae. Finally, we used multiple genetic techniques to reveal new information about the physiology of L. mimadelfi. We find no evidence for the presence of hindgut parabasalians, but we do find evidence of indirect wood-feeding – xylomycophagy. Metabarcoding of gut contents shows that L. mimadelfi is a generalist feeder, but wood-associated fungi are the most consistent and abundant food across samples, comprising about 40% of its diet. We then discuss all the results to hypothesize possible evolutionary sequences for the acquisition of direct wood-feeding in Kittrickea, and the shift from generalist to specialist feeding.

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DOI: 10.3897/asp.84.e185452

Authors: Qi Feng, Xisen Jia, Chenxin Zhou, Dong Ren, Yongjie Wang

Abstract: Abstract Therevidae is a diverse family of lower Brachycera, yet its early evolutionary history remains poorly understood due to the absence of available Mesozoic fossils. Here, we describe †Paleothereva longicoxa gen. et sp. nov. from mid-Cretaceous Kachin amber, representing the oldest member of known derived therevids. Phylogenetic analysis and morphological comparisons integrating both extant and fossil species, consistently place this fossil within Taenogera genus-group of Agapophytinae. This finding indicates that Agapophytinae had already diverged by the mid-Cretaceous, suggesting an earlier origin of Therevidae than previously inferred from molecular data. The occurrence of Paleothereva on the Gondwanan-derived West Burma Block further supports the hypothesis that the ancestors of Agapophytinae—or possibly the broader Agapophytinae + Therevinae clade—originated in Gondwana. Ancestral character state reconstruction of female terminalia within therevid flies reveals that Paleothereva and extant relatives likely shared similar oviposition behaviors associating with sandy-soil environments, which may have driven the corresponding specialization and diversification of acanthophorite spines in female. These new findings provide rare, direct evidence illuminating the early evolution of derived therevid lineages.

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DOI: 10.3897/asp.84.e182696

Authors: Leonela Olivera, María Cecilia Melo, Pablo Matías Dellapé

Abstract: Abstract Acanthocephala, with 40 known species, is the most diverse genus of the tribe Acanthocephalini, which contains some of the largest species of Coreidae. In this contribution, Acanthocephala guatemalena, A. granulosa, A. luctuosa, A. panamensis, and A. thoracica were resurrected as valid species, and a morphological phylogenetic analysis, using continuous and discrete characters, was performed to test the monophyly of the genus. The analysis was carried out under implied weighting, including 34 species of Acanthocephala as ingroup, and 20 species from other genera of Acanthocephalini and Placoscelini as outgroups. Acanthocephala was recovered as a monophyletic group, and sister of the clade (Spilopleura parensis + S. ochracea). To formalize and show the relationships of the species in Acanthocephala, the subgenera A. (Acanthocephala) and A. (Metapodiessa) were revalidated, and three new subgenera were proposed: A. (Spinipedia) subgen. nov., A. (Contrastata) subgen. nov., and A. (Pronoptera) subgen. nov. Additionally, a detailed redescription of Acanthocephala, a subgeneric key, descriptions and redescriptions for all subgenera, photographs of dorsal and ventral habitus of the type species of each subgenus, and photographs of the diagnostic characters, including male and female genitalia are provided.

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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.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.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.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.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.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.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.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.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.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.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.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.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.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.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.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.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.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.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.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.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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DOI: 10.3897/asp.81.e102683

Authors: Ya-Lian Wang, Zi-Qi Peng, Feng-Liang Jin, Hong-Wei Chen, Jia Huang

Abstract: With the increasing number of cryptic species discovered in major branches of the subfamily Steganinae, accurate species identification is becoming challenging. Possible evolutionary stasis often gives rise to the formation of morphological conservatism and cryptic species, as is the case for the members of the Stegana (subgen. Orthostegana). In this study, the species boundaries of all Asian Orthostegana were accessed based on two mitochondrial genes [cytochrome c oxidase subunit I (COI) and NADH dehydrogenase subunit 2 (ND2)] using different species delimitation methods and evolutionary models. By integrating morphological and molecular evidence, we clearly validated the evolutionary independence of all five known species and identified nine new species from Southwest China: Stegana (Orthostegana) aini Peng & Chen sp. nov., S. (O.) brevivittata Peng & Chen sp. nov., S. (O.) cuodi Peng & Chen sp. nov., S. (O.) fuscofemorata Peng & Chen sp. nov., S. (O.) latipalpula Peng & Chen sp. nov., S. (O.) macrostephana Peng & Chen sp. nov., S. (O.) mohnihei Peng & Chen sp. nov., S. (O.) obscurala Peng & Chen sp. nov., and S. (O.) pinguitia Peng & Chen sp. nov. Moreover, we discussed the relationship between the Asian Orthostegana lineage and the subgen. Oxyphortica, and the necessity of taxonomic re-evaluation for the Neotropical species with molecular data. The high morphological conservatism within Orthostegana is largely the outcome of in situ diversification for adapting to the ecological environment.

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DOI: 10.3897/asp.81.e100520

Authors: Leif Moritz, Antonio Parra-Gómez

Abstract: The millipede family Siphonorhinidae (order Siphonophorida) shows a scattered distribution in South Africa, Madagascar, India, Southeast Asia, and North America. So far, the family is unknown from South America, while species of Siphonophoridae, the second family of the order, are relatively abundant on the continent. However, not a single Siphonophorida is known from Chile. Here we describe the monotypic genus Notorhinus gen. nov. with N. floresi sp. nov. and record a second Notorhinus (undescribed) species, as first records of the order Siphonophorida in Chile and of the family Siphonorhinidae in South America. Notorhinus gen. nov. is distinct from the remaining Siphonorhinidae by the arrangement of the sensilla basiconica on the antennae and other somatic and sexual characters. However, it shows close morphological affinities to the North American genus Illacme Cook and Loomis 1928. In the Americas Siphonorhinidae were previously only known from California (USA), where they inhabit subterranean micro-habitats. The Chilean species was found under a piece of decaying wood in a small patch of fragmented native forest. Thus, the group shows a disjunct antitropical distribution in America at ca. 37° North and 38° South. They might be the relict of a once greater distribution, which persisted in these areas due to similar climatic conditions.

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DOI: 10.3897/asp.81.e100385

Authors: Martin Fikáček, Jan Simon Pražák, Andrew E. Z. Short, François Rion

Abstract: We describe the first definite fossil of the water scavenger beetle subfamily Enochrinae (Coleoptera: Hydrophilidae): Cymbiodyta samueli sp. n. from the Eocene Baltic amber from the Lithuanian coast. The new species is extremely similar and likely closely related to the only European species, C. marginella and confirms the European occurrence of the genus since the Eocene. A reanalysis of the historical biogeography of the genus, including the fossil taxon, revealed a wide Euro-American distribution of the ancestor of all modern species of the genus, corresponding to the position of landmasses and existing land connections between North America and Europe in the Late Cretaceous. The biogeographic reconstructions and the fossil both suggest that European Cymbiodyta is an ancient relict lineage which used to be more diverse in the past but survived until today in a single species C. marginella.

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DOI: 10.3897/asp.81.e89641

Authors: Leon Gustavo de Miranda Tavares, Renato Jose Pires Machado, Adolfo Ricardo Calor

Abstract: A taxonomic revision and the first phylogenetic hypothesis of the Neotropical genus Ameromyia Banks (Myrmeleontidae: Brachynemurini) is herein presented. The phylogeny is based on 45 morphological characters and recovered the traditional Ameromyia as paraphyletic in respect to the monotypic genus Venezueleon Stange, which is here synonymized under Ameromyia. Three species are synonymized (A. hirsuta Navás and A. stevensi Navás under A. nigriventris (Walker), and A. pentheri Navás under A. strigosa (Banks)) and two new species are described (A. clepsydra sp. nov. and A. explicata sp. nov.). Ameromyia sensu novo is a valid genus with 12 species restricted to South America, and divided into two species groups. Taxonomic keys are also presented to adults and larvae, as well as a discussion on the genus biology.

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DOI: 10.3897/asp.81.e101441

Authors: Jindřich Roháček, Jörg U. Hammel, Viktor Baranov

Abstract: A new family of Diptera Acalyptratae, Christelenkidae Roháček fam. nov., is established for Christelenka multiplex Roháček gen. et sp. nov., an unusual extinct taxon described from a unique male specimen preserved in Baltic amber (Mid-late Eocene, ca 48–34 Ma). Apart from detailed examination by light microscopy and photography, the holotype of the new species has also been studied by means of X-ray synchrotron microtomography with the aim of obtaining additional morphological data for consideration of its relationships. Because of a very peculiar combination of morphological characters, the new family is tentatively considered a separate lineage of Acalyptratae having no apparent sister-group relationship with any of the known families. Its probable relationships to some families of Opomyzoidea and Ephydroidea are discussed.

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DOI: 10.3897/asp.81.e96397

Authors: Augusto Henrique Batista Rosa, Eduardo de Proença Barbosa, Patrícia Avelino Machado, Ricardo Russo Siewert, André Victor Lucci Freitas

Abstract: Dasyophthalma includes five species of medium-sized butterflies, all endemic from the Atlantic Forest of Brazil. All known species are univoltine and are dayflying, differently from other Brassolini that are mostly crepuscular. In despite of recent advances little is known about their natural history. Three out of the five species are included in the Brazilian Red List of threatened fauna and are of conservation concern. The present study provides for the first time a phylogenetic assessment of all Dasyophthalma species based on a molecular approach based on three loci. Also, the taxonomic status of D. rusina delanira was revised based on molecular data. In addition, up-to-date distributional data and conservation aspects of the threatened species from the genus are presented and discussed. The molecular phylogenetic analysis supports the monophyly of Dasyophthalma, with Dynastor darius as its sister-group, and, combined with a genetic divergence analysis, supported Dasyophthalma delanira stat. rest. as a valid name to species-level, sister-group to D. geraensis (and not a subspecies of D. rusina). The geographical range (extent of occurrence and area of occupancy) for all five species are presented, showing that these are very restricted for D. delanira stat. rest. and D. geraensis, following the distributions of the high-altitude forests. As much biological information about the genus is lacking, the present study can serve as a starting point for future studies on Dasyophthalma, adding information that can be crucial for future conservation actions and essential to assure the future of the threatened species in this genus.

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DOI: 10.3897/asp.81.e102404

Authors: Jan Simon Pražák, Martin Fikáček, Jakub Prokop, Tomáš Lackner

Abstract: We describe three new genera and four new species of the Histeridae (Coleoptera) from the mid-Cretaceous amber in Myanmar. Platycretus muscularis Simon Pražák & Lackner gen. & sp. nov. represents the first known fossil of the subfamily Histerinae from the Cretaceous. We assign the remaining three fossils, Olexum complanatum Simon Pražák & Lackner gen. & sp. nov., Cretanapleus seideli Simon Pražák & Lackner gen. & sp. nov., and Yethiha pubescens Simon Pražák & Lackner sp. nov. to the subfamily Dendrophilinae. Platycretus muscularis and O. complanatum have adaptations typical for the subcortical lifestyle (flattened body shape, dilated protibiae), proving this life strategy existed in independent lineages of clown beetles already in the Cretaceous. We also provide a review of all Histeridae fossil species described up to date and test the phylogenetic position of all of them including the newly described ones.

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DOI: 10.3897/asp.81.e85571

Authors: Andrés F. García, Osvaldo Villarreal

Abstract: A morphological cladistic analysis, consisting of 45 terminals and 58 characters was performed to evaluate the monophyly of the genus Paravima Caporiacco, 1951, and its relationship to other Leiosteninae. The analysis resulted in two most parsimonious trees, all recovering Paravima as monophyletic with the inclusion of two species formerly described in Avima Roewer, 1949, therefore, the following nomenclatural acts are herein proposed: Paravima plana (Goodnight & Goodnight, 1949) comb. nov., and Paravima quirozi (González-Sponga, 1981) comb. nov., Avima vigirima (Villarreal-M & Rodríguez-Manzanilla, 2003) is here considered as a junior subjective synonym of P. quirozi (González-Sponga, 1981) comb. nov., Paravima acanthoconus Villarreal-Manzanilla & DoNascimiento, 2005 is here considered as a junior subjective synonym of Paravima propespelunca González-Sponga, 1987, and Paravima flumencaurimarensis González-Sponga, 1987 is here considered as a junior subjective synonym of Paravima goodnightorum Caporiacco, 1951. Additionally, three new species are described: P. lokura sp. nov. (Tamá National Natural Park, Norte de Santander department) and P. magistri sp. nov. (Los Tunos Natural Reserve, Cundinamarca department), both from Colombia; and P. totoro sp. nov. (Henri Pittier National Park, Aragua state) from Venezuela. Finally, an emended generic diagnosis, a key for all the species, and a distributional map are presented.

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DOI: 10.3897/asp.81.e96319

Authors: Anna Paśnik, Sebastian Tarcz, Łukasz Przybyłowicz

Abstract: The subgenus Parapisa of the genus Apisa is reviewed based on the examination of 104 specimens. Apisa (P.) cinereocostata and A. (P.) subargentea are redescribed and their intraspecific variation is analysed in detail. A new species A. (P.) asipa, similar in the general coloration to other Apisa taxa, but very distinctive in the male genital morphology and the shape of the wing scales, is described from Cameroon and Nigeria. Apisa (P.) cinereocostata is hypothesized to be a widespread, but highly polymorphic taxon with significant variation in body size, intensity of grey coloration, and the proportions and shape of certain morphostructures of male genitalia. Determination keys and extensive illustrations of the variation are provided to enable proper identification of specimens.

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DOI: 10.3897/asp.81.e96737

Authors: Manuel Brazidec, Frédéric Legendre, Vincent Perrichot

Abstract: The Lancepyrinae are an extinct subfamily of Bethylidae known exclusively from Cretaceous amber deposits of Lebanon, Spain, Taimyr and Myanmar. In this study, we describe and illustrate four new genera and five new species of lancepyrine wasps from the Albian of Hkamti and late Albian-early Cenomanian of Kachin (Myanmar): Azepyris delamarrei gen. et sp. nov., Burmapyris ohmkuhnlei sp. nov., Gwesped groehni gen. et sp. nov., Paralanceis chotardi gen. et sp. nov. and Yunbayin rossei gen. et sp. nov. These taxa not only highlight the taxonomic diversity of the Lancepyrinae during the mid-Cretaceous but they also reveal the morphological disparity of the subfamily. To establish the phylogenetic relationships of these fossils and to check the monophyly of the Lancepyrinae, we add them to a pre-existing morphological matrix and perform a cladistic analysis. We retrieve the subfamily as poorly supported yet monophyletic, with the newly described taxa deeply nested in it. A key to the genera of Lancepyrinae is provided. Finally, we erect the subfamily Cretabythinae subfam. nov. for the genera Cretabythus Evans, 1973, Holopsenelliscus Engel, 2019 and Megalopsenella Jouault et al., 2020 as no taxonomic treatment has been provided for these taxa after the transfer of Holopsenella Engel et al., 2016 as Aculeata incertae sedis.

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DOI: 10.3897/asp.81.e95181

Authors: Ricardo Botero-Trujillo, Leonel Martínez, Hernán Augusto Iuri, Andrés Alejandro Ojanguren-Affilastro, Leonardo Sousa Carvalho

Abstract: The solifuge genera Eutrecha Maury, 1982 and Xenotrecha Maury, 1982 are among the rarest and most elusive groups of camel spiders in the New World. Since their inception, both genera have remained unclassified within the subfamilial scheme of Ammotrechidae, where they belong, and their systematic position and affinities to other genera continue to be unexplored. This contribution addresses the affinities that Eutrecha and Xenotrecha have to the type genus of Ammotrechinae, Ammotrecha Banks, 1900. Based on the taxonomic distribution of characters shared by these genera, it is proposed that the three are closely related and classified into Ammotrechinae. Revised diagnoses are presented for Ammotrechinae s.str., Eutrecha, and Xenotrecha, and redescriptions are presented for species of these genera, when possible. A new species, Eutrecha belenensis sp. nov., is described from Colombia, raising to three the number of species in the genus. New material of Eutrecha florezi Villareal-Blanco, Armas and Martínez, 2017 and Xenotrecha huebneri (Kraepelin, 1899) is referenced, thus extending the distribution range of these species in Colombia and Brazil, respectively. Ammotrechinae s.str. is here defined by the presence of a retroventral longitudinal carina on the movable finger of the chelicerae and a cleavage plane basally on the femur of pedipalps, among other characters. The cleavage plane allows the pedipalp to be autotomized, representing the first report of pedipalp autotomy in Solifugae. This work also presents a discussion on the taxonomy of Ammotrechidae and delves into some aspects that affect the current delimitation of some of its subfamilies.

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DOI: 10.3897/asp.81.e98418

Authors: Jiří Kolibáč, Kateřina Rosová, Jan Simon Pražák, Jörg U. Hammel, Jakub Prokop

Abstract: A larva of Cleroidea in Burmese amber is described, which is the first record of an immature beetle of the basal cucujiform superfamily for the Mesozoic. Well-preserved unique specimen is described and illustrated using traditional methods as well as synchrotron-radiation-based micro-computed tomography (SRµCT) to reconstruct the specimen and discern integumental details of cephalic structures, especially the mouthparts. Cretorhadalus constantini gen. et sp. nov. is unambiguously assigned to the melyrid lineage of Cleroidea and tentatively classified within the basal family Rhadalidae. Within this family, this fossil larva has the ancestral cleroid pattern of the stemmata (2+3) and well-developed hooked urogomphi. Based on a comparison with extant rhadalids, as well as most members of the melyrid lineage, the larvae and adults of this new species were probably carnivorous, living on the trunks and branches of trees or in galleries where they foraged for soft xylophagous insects.

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DOI: 10.3897/asp.81.e85191

Authors: Si Xun Ge, Zhuo Heng Jiang, Jia Qi Wang, Kui Song, Chao Zhang, Shao Ji Hu

Abstract: The taxonomic status of the Pieris napi-complex and similar species which occur in China are revised. Relevant species distributed in the adjacent regions were included to clarify the status of Chinese species and were briefly revised. All those species are described and illustrated and new synonyms are established. A molecular phylogenetic analysis is also performed on the species group including similar species, to investigate the phylogenetic relationships between taxa. Species of the Pieris napi-complex that occur in China and adjacent regions are redefined, with four similar species excluded (P. melaina, P. extensa, P. chumbiensis gyantsensis and P. melete). A distribution map and keys of the complex including similar species are provided. The taxon P. mihon Yakovlev, 2006 stat. nov. is raised from subspecies to species status; P. narina Verity, 1908 stat. rev. is confirmed as a distinct species rather than a subspecies of P. ochsenheimeri; Pieris euorientis Verity, 1908 stat. rev. is recovered as a distinct species sister to P. dulcinea. Two taxa, ssp. sauron and ssp. bryonides are moved from subspecies of P. euorientis and P. bryoniae, respectively, to P. napi, i.e. P. napi sauron Yakovlev, 2004 comb. nov and P. napi bryonides Sheljuzhko, 1910 comb. rev. A new synonym is proposed: Pieris ochsenheimeri tianshansis Tadokoro, Shinkawa & Wang, 2014, new synonym of P. mihon Yakovlev, 2006. A new mistaken identification is proposed: Pieris dulcinea kneitzi is a misidentification of Pieris erutae kneitzi Eitschberger, 1983 comb. rev. Five Chinese species belonging to the Pieris napi-complex were confirmed, namely P. narina, P. mihon, P. latouchei, P. dulcinea, and P. erutae. Among them, two species, P. mihon Yakovlev, 2006 and Pieris narina Verity, 1908, are newly recorded from China. The taxonomic status of Pieris steinigeri Eitschberger, 1983 and Pieris bryoniae sifanica Grum-Grshimailo, 1895 is also discussed.

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DOI: 10.3897/asp.81.e93570

Authors: Ante Vujić, Snežana Radenković, Anatolij Barkalov, Nataša Kočiš Tubić, Laura Likov, Tamara Tot, Grigory Popov, Alex Prokhorov, Ebrahim Gilasian, Shehzad Anjum, Mihajla Djan, Banafsha Kakar, Andrijana Andrić

Abstract: The phytophagous hoverfly genus Merodon Meigen, 1803 (Diptera, Syrphidae) presently comprises 205 described species distributed in the Palaearctic and Afrotropical regions, and it is differentiated into multiple species groups. In this work, we revise the Merodon tarsatus species group, providing an illustrated key to species, a detailed discussion on relevant taxonomic characters, and a morphological diagnosis. We summarize morphological characteristics of the 12 species in the M. tarsatus group, together with available distributional data. Moreover, we provide descriptions for five new species, namely Merodon angustitarsis Vujić & Gilasian sp. nov., M. dumosus Vujić, Likov & Radenković sp. nov., M. latiantennatus Vujić, Popov & Prokhorov sp. nov., M. namaghijamii Vujić, Likov & Radenković sp. nov. and M. pakistanicus Vujić, Likov & Radenković sp. nov. Additionally, we describe one more species, Merodon nigroalatus Vujić, Likov & Radenković sp. nov., that belongs to the same lineage and is related to the Merodon tarsatus group based on molecular markers and structure of male genitalia, but lacks modifications of the metatarsus and sternum 4 that are the main diagnostic features of the group. Furthermore, we redefine and redescribe seven taxa of the M. tarsatus group. Following detailed analyses of type material, we revise the status of all available taxa related to M. tarsatus and we propose the following new synonyms: M. ankylogaster Hurkmans, 1993 syn. nov. (junior synonym of M. hypochrysos Hurkmans, 1993); M. persicus Hurkmans, 1993 syn. nov. (junior synonym of M. oidipous Hurkmans, 1993); and M. xanthipous Hurkmans, 1993 syn. nov. (junior synonym of M. marginicornis Hurkmans, 1993). We apply Maximum Parsimony and Maximum Likelihood analyses of the 5′ end of the mitochondrial COI gene sequences to genetically characterize species in the M. tarsatus group and the related species, M. nigroalatus Vujić, Likov & Radenković sp. nov.

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DOI: 10.3897/asp.81.e86898

Authors: Ayman Khamis Elsayed, Tadao Ichita, Makoto Tokuda

Abstract: We describe a gall midge Parampelomyia yukawai Elsayed and Tokuda gen. nov. sp. nov. belonging to the subtribe Schizomyiina (Diptera: Cecidomyiidae: Asphondyliini) based on an integrative taxonomic study. This species forms barely-swollen flower bud galls on the porcelain berry Ampelopsis brevipedunculata var. heterophylla (Vitaceae) in Japan. The new genus is distinguishable from all known schizomyiine genera by tarsomere I lacking a ventroapical extension, the bulbous base of the protrusible portion of the ovipositor, the fused and sclerotized female cerci, the bidentate gonostylus, and the larval terminal abdominal segment that bears two corniform, two asetose and six setose papillae. The new genus is compared with and separated from the similar genera Schizomyia and Ampelomyia morphologically and phylogenetically.

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DOI: 10.3897/asp.81.e81961

Authors: Alexandre Cruz Domahovski, Rodney Ramiro Cavichioli

Abstract: Based on a phylogenetic analysis using implied weighting for 59 taxa and 155 morphological characters, we provided a taxonomic revision for Acuera. The genus was recovered as polyphyletic; herein we described three new genera and 16 new species, and proposed 23 new combinations, a new status and four new synonyms for Gyponini. Coarctana gen. nov. is described to include 15 species: C. rana (DeLong & Freytag) comb. nov. (type species) transferred from Acuera; five species transferred from Curtara, C. apena (DeLong & Freytag) comb. nov., C. pedica (DeLong) comb. nov., C. punctata (Spångberg) comb. nov., C. secara (DeLong & Triplehorn) comb. nov., and C. serpenta (DeLong) comb. nov.; and nine new species, C. asymmetrica sp. nov., C. glabra sp. nov., C. nigromedia sp. nov., C. occultata sp. nov., C. oricula sp. nov., C. parvula sp. nov., C. striata sp. nov., C. taurina sp. nov., and C. vilavelha sp. nov. Effossana gen. nov. includes eight species: two species transferred from Acuera, E. gloma (DeLong & Freytag) comb. nov. (type species) and E. fructa (DeLong & Freytag) comb. nov.; two species transferred from Curtara, E. gatuna (DeLong & Wolda) comb. nov. and E. scapa (DeLong) comb. nov.; and four new species, E. assimulata sp. nov., E. circumnota sp. nov., E. intrinseca sp. nov., and E. tenuemarginata sp. nov. A new monotypic genus, Propincurtara gen. nov., is described to include P. longilinea sp. nov. The subgenus Parcana is raised to genus rank, comprising nine species: P. atitlana (Fowler) comb. nov., P. concilia (DeLong & Freytag) comb. nov., P. extara (DeLong & Freytag) comb. nov., P. mimica (DeLong & Freytag) comb. nov., P. spreta (Fowler) comb. nov., and P. ultima (DeLong & Freytag) comb. nov. (type species); also three species transferred from Curtara, P. albororata (Fowler) comb. nov., P. animosa (DeLong & Freytag) comb. nov., and P. ventusa (DeLong & Freytag) comb. nov. Curtara lineatana DeLong & Freytag is a junior synonym of P. concilia (DeLong & Freytag) comb. nov., Acuera inlustra DeLong & Freytag is a junior synonym of Hamana manifesta DeLong, and A. culmena DeLong & Freytag and A. prodiga DeLong & Freytag are synonyms of A. menaca DeLong & Freytag. Four species of Acuera are transferred to Curtara: C. facera (DeLong & Freytag) comb. nov., C. labella (Osborn) comb. nov., C. nama (DeLong & Freytag) comb. nov., and C. patula (DeLong & Freytag) comb. nov. Two new species, A. extenuata sp. nov. and A. umbra sp. nov. are described and placed in the subgenus Acuera. We provide descriptions and illustrations of the new taxa, redescription of the type species of the genera Acuera, Coarctana gen. nov., Effossana gen. nov., and Parcana stat. nov., 40 new country and Brazilian states records, and key to males of each genus.

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DOI: 10.3897/asp.81.e85755

Authors: Tatiana Petersen Ruschel, Filipe Michels Bianchi, Luiz Alexandre Campos, Gervásio Silva Carvalho

Abstract: The Linnean, Wallacean, and Darwinian shortfalls are knowledge gaps about species taxonomy, distribution, and evolution, respectively. Fidicinini is a tribe of cicadas that suffers from these gaps. We assessed specimens of the tribe sharing similar male genital shape (uncus), but fitting the somatic morphology of either Dorisiana Metcalf, 1952 and Guyalna Boulard & Martinelli, 1996. We build a phylogenetic hypothesis by total evidence analysis and perform a character optimization of the uncus and timbal cover shapes, the last used as diagnostic for both genera. Dorisiana and Guyalna were recovered non-monophyletic. A new genus, Acanthoventris gen. nov., and ten new species are proposed: A. charrua Ruschel sp. nov., A. claudiae Ruschel sp. nov., A. faustopsaltrius Ruschel sp. nov., A. iara Ruschel sp. nov., A. igneus Ruschel sp. nov., A. olivarius Ruschel sp. nov., A. phoenix Ruschel sp. nov., A. rubemi Ruschel sp. nov., A. tumidus Ruschel sp. nov., and A. viridinotatus Ruschel sp. nov.; and three new combinations A. densusus (Boulard & Martinelli, 2011) comb. nov., A. drewseni (Stål, 1854) comb. nov., and A. jauffreti (Boulard & Martinelli, 2001) comb. nov. We provide illustrated identification keys, occurrence maps, and discuss the distribution of the species in the new genus.

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