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.e178495

Authors: Wen-Jun Cao, Lu Jiang

Abstract: Abstract Larvae of the megadiverse Scarabaeoidea play important economic and ecological roles, and their mouthparts provide valuable morphological clues for species identification and dietary inference. However, the underlying relationships between morphological characters and feeding habits remain insufficiently elucidated. In this study, the larval mouthparts of 22 species representing ten major lineages of Scarabaeoidea were examined in detail. Seventeen characters of the larval mouthparts were analyzed within a phylogenetic framework. Three characters were identified as candidate synapomorphies for the major lineages represented here: maxillary stridulatory teeth for Scarabaeidae, mandibular stridulatory ridges for Cetoniinae + (Dynastinae + Rutelinae), and a circular arrangement of epipharyngeal phobae for Aphodiinae + Scarabaeinae. By contrast, several homoplastic mouthpart traits were recurrently associated with larval feeding habits in the sampled taxa, including a setose acroparia, heli on the epipharynx, and a blade-like incisor region in phytophagous lineages, and the circular phoba complex together with the absence of nesium in coprophagous lineages. These results suggest that larval mouthparts in Scarabaeoidea may retain both phylogenetic and trophic signals, providing a morphological basis for interpreting the evolution of feeding habits in scarab larvae.

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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.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.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.e151555

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

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

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DOI: 10.3897/asp.83.e150736

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

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

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DOI: 10.3897/asp.83.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.82.e132546

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

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

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DOI: 10.3897/asp.82.e106391

Authors: Chong Li, Liang Tang

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

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DOI: 10.3897/asp.82.e131738

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

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

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DOI: 10.3897/asp.82.e112684

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

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

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DOI: 10.3897/asp.82.e114508

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

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

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DOI: 10.3897/asp.81.e98166

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

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

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DOI: 10.3897/asp.81.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.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.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.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.e101411

Authors: Wei Zhao, Hao Yu Liu, Xue Ying Ge, Yu Xia Yang

Abstract: The resolution of phylogenetic relationship among animals is still one of the most challenging problems in systematic zoology. Insect wing is a highly valued morphological character in the systematics, but few studies have been conducted to quantify wing shape variations for phylogenetic reconstruction. In this study, with Cantharinae as the subject, we conducted the GM analyses from hindwings of 16 representative genera. Further, we conducted the UPGMA based on Procrustes distance and Euclidean similarity measure of Mahalanobis distance, respectively, and NJ analysis of the Mahalanobis distance, as well as MP analysis using merged landmark dataset. In the meantime, we constructed the phylogenetic relationships among these genera based on the mitochondrial genomes, with a total of 41 sequences novel to Cantharinae, by BI and ML analyses. As a result, the CVA analysis demonstrated that the hindwing shapes of the cantharid genera are significantly different from one another. All the topologies produced by the GM data partially correspond with that of mitogenomic data. The close relationships of some genera are frequently recovered, including Cyrebion + Themus, Cantharis + Taiwanocantharis + Taocantharis, Stenothemus + Falsopodabrus + Habronychus. These results prove the importance and potential application of the hindwing shapes in recovering the relationships among the sibling genera.

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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.80.e84013

Authors: Marek Linský, Zuzana Čiamporová-Zaťovičová, Kristína Laššová, Fedor Čiampor Jr

Abstract: The riffle beetle genus Hexanchorus Sharp is, with 25 known species, the most speciose genus of the subfamily Larainae in the Neotropics and the second largest globally. An analysis of its phylogeny, based on two mitochondrial (COI, 16S) and two nuclear (18S, 28S) markers, including Hexanchorus-like, but morphologically distinct specimens, supported presence of an unknown genus. The new genus, Rumilara gen. nov., is described here with four new species (R. obscura sp. nov., R. paterna sp. nov., R. riberai sp. nov., R. suppressa sp. nov.) and their larvae. The separate position of the new genus is, beside molecular differences, well substantiated by the morphology of adults (the absence of sexual dimorphism, structure of the pronotum) and larvae (pleurites never reaching to sixth abdominal ventrite). H. sagittatus stat. nov. is elevated to species rank based on molecular data, and Rumilara leleupi (Delève) comb. nov. is transferred from the genus Hexanchorus and redescribed, based on its morphology.

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DOI: 10.3897/asp.80.e84114

Authors: Jan Erik Sedlmeier, Arnaud Faille

Abstract: The past climatic changes caused repeated distribution shifts within insect populations leading to a highly diverse fauna in the mountain regions, which have acted as a refuge for many groups. There, some taxa have adapted to high altitudes and cold climatic conditions. One of those is the highly diverse and Holarctic subgenus Cryobius Chaudoir, 1838 (Carabidae: Pterostichus) including both locally and widely distributed species. Isolated and morphologically divergent populations of the same species led to the description of many subspecies. Until now, there has been no comprehensive work concerning the phylogeny of Cryobius, and genetic data on this taxon are sparse. This study is the first to provide insights into the molecular phylogeny of this subgenus, focusing on species from the Pyrenean and Cantabrian mountain systems. Cryobius specimens were sequenced targeting mitochondrial and nuclear genes. A molecular phylogeny was then built, merging the new data with genetic data from online public databases. All species of Cryobius included in this study form a monophyletic clade within Pterostichus. The synonymy of the two former taxa Pyreneorites and Haptoderus with Cryobius is confirmed by this study. Cryobius of the Pyreneo-Cantabrian area are closely related. Moreover, several well-supported clades of local species were found. The results further indicate a relation between Nearctic and Eastern Palearctic Cryobius, in agreement with the theory of faunal and floral colonization of North America via the Bering land bridge.

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DOI: 10.3897/asp.80.e86582

Authors: Mario Schädel, Margarita Yavorskaya, Rolf Beutel

Abstract: The earliest known fossil beetle †Coleopsis archaica is re-examined using Reflectance Transformation Imaging (RTI). The morphological observations are evaluated with respect to phylogenetic implications and the early evolution of Coleoptera. †Coleopsis archaica belongs to an early Permian branch of beetles, outside a monophyletic unit comprising Coleoptera (in the widest sense) excluding †Tshekardocoleidae. This clade is mainly characterized by a complex of apomorphic features: elytra with epipleura and with a close fit with the posterior body, thus forming a tightly sealed subelytral space. In contrast to this, the elytra of †C. archaica and †Tshekardocoleidae cover the metathorax and abdomen in a loose tent-like manner and posteriorly distinctly surpass the abdominal apex. So far, no synapomorphies of the two taxa from the first half of the Permian have been identified. The very short and transverse pronotum is likely an autapomorphy of †C. archaica. A thorough documentation of the structural features of early beetle fossils should have high priority. RTI is a very promising tool to obtain new and well-founded morphological data, which will allow a thorough phylogenetic evaluation of Permian beetles in future studies. We extended the conventional RTI workflow by focus merging and panoramic stitching, in order to overcome previous limitations. Taxonomic re-arrangements of stem group beetles including †C. archaica were suggested in recent studies by A.G. Kirejtshuk and co-workers. As they are not based on shared derived features they are irrelevant in a phylogenetic and evolutionary context.

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DOI: 10.3897/asp.80.e81736

Authors: Yan-Da Li, Yu-Bo Zhang, Karol Szawaryn, Di-Ying Huang, Chen-Yang Cai

Abstract: The family Corylophidae is a moderately diverse coccinelloid beetle family. The fossil record of corylophid beetles is extremely sparse, with only one species formally described from the Eocene Baltic amber. Here we report a new corylophid genus and species, Xenostanus jiangkuni Li, Szawaryn & Cai gen. et sp. nov., from mid-Cretaceous amber from northern Myanmar (ca. 99 Ma). Xenostanus is most distinctly characterized by the antenna with 10 antennomeres and the presence of metaventral and abdominal postcoxal lines. Our phylogenetic analysis suggested Xenostanus as sister to tribe Stanini. Based on its distinctive morphology and the phylogenetic results, Xenostanus is placed in the tribe Xenostanini Li, Szawaryn & Cai trib. nov.

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DOI: 10.3897/asp.80.e82644

Authors: Zi-Wei Yin, Erik Tihelka, Jesus Lozano-Fernandez, Chen-Yang Cai

Abstract: The extant tropical tribe Hybocephalini is a morphologically highly derived group of the subfamily Pselaphinae (Coleoptera: Staphylinidae), which is characterized most notably by the modified squamous setae that cover various parts of the body. Ten genera and 69 extant species have been found in the Afrotropical and Oriental regions, with one species found in northern Australia. Prior to this study the evolutionary history of the tribe has been remained elusive due to the dearth of known fossils. Here, we describe the first fossil representative of Hybocephalini, Europharinodes schaufussi Yin & Cai gen. et sp. nov., based on an adult male preserved in Baltic amber (ca 45.0–38.0 Ma). Using X-ray microtomography, the anatomy including the endoskeletal structures of the head, the full pattern of foveation, and the aedeagus of the beetle were reconstructed. Europharinodes shares most derived traits that are congruent with extant members of Hybocephalini, but it also possesses plesiomorphic and autapomorphic characters unknown in living relatives. In order to constrain the phylogenetic placement of Europharinodes, we created an updated morphological character matrix to explore relationships among this genus and related groups. A monophyletic Hybocephalini was recovered by maximum likelihood and parsimony analyses, with Europharinodes being well-resolved as sister to all modern relatives in the likelihood tree. The fossil thus sheds new light on the morphological evolution of Hybocephalini and suggests a broader palaeodistribution of the tribe during the middle Eocene. The disjunct distribution of an Eocene Baltic amber species and an extant Afrotropical-Oriental distribution of the tribe is probably relictual, and was shaped by global cooling during the Eocene–Oligocene transition.

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DOI: 10.3897/asp.80.e76575

Authors: André Silva Roza, Carlos G. Schrago, José Ricardo M. Mermudes

Abstract: In this study, we aim to understand the boundaries and the species evolutionary relationships in Mesochila, which includes 20 species arranged into three subgenera. We also conducted a biogeographic analysis that allowed us to identify the major events that shaped the currently disjunct distribution of the group. Our analyses were performed employing four major phylogenetic algorithms: equal and implied weight parsimony, maximum likelihood and Bayesian inference. Phylogenetic analyses, including 25 taxa (all 20 species in the genus plus five outgroups) and 37 characters, indicated that Mesochila is a monophyletic group composed of four strongly supported lineages, although no consensus was attained regarding relationships between lineages. Bayesian inference resulted in the less resolved topology, whereas implied weight parsimony yielded the most different tree when using a low k value (1). We present a new infrageneric classification for the group with the description of Mesochila (Neomesochila) subgen. nov. to accommodate M. (N.) brevipennis, M. (N.) drechseli, M. (N.) moraveci, and M. (N.) prepusula. Furthermore, we transfer M. (M.) distincta from M. (Eumesochila) back to M. (Mesochila). The biogeographical analysis suggested a South American origin for the group, in Chacoan/Paraná dominions (Atlantic rainforest/Cerrado biomes), with subsequent dispersions to Central America and Amazonia.

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DOI: 10.3897/asp.79.e72724

Authors: Yan-Da Li, Erik Tihelka, Zhen-Hua Liu, Di-Ying Huang, Chen‑Yang Cai

Abstract: Abstract The cryptic slime mold beetles, Sphindidae, are a moderately diverse cucujoid beetle family, whose members are obligately tied to slime molds throughout their life. The fossil record of sphindid beetles is sparse; stem-sphindids and crown-group members of uncertain systematic placement have been reported from Cretaceous ambers. Here we review the Mesozoic fossil record of Sphindidae and report a new sphindid genus and species, Trematosphindus newtoni gen. et sp. nov., from Albian/Cenomanian amber from northern Myanmar (ca. 99 Ma). Trematosphindus is set apart from all other sphindids by the presence of distinct lateral cavities on the anterior pronotal angles. Our phylogenetic analysis identifies Trematosphindus as an early-diverging genus within Sphindidae, sister to the remainder of the family except Protosphindus, or Protosphindus and Odontosphindus. The new fossils provide evidence that basal crown slime mold beetles begun to diversify by the mid-Cretaceous, providing a valuable calibration point for understanding timescale of sphindid co-evolution with slime molds.

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DOI: 10.3897/asp.79.e65325

Authors: Lech Karpiński, Wojciech T. Szczepański, Radosław Plewa, Lech Kruszelnicki, Katarzyna Koszela, Jacek Hilszczanski

Abstract: This paper sheds the first light on the phylogeny of the Central Asian genus Turanium Baeckmann, 1922. By applying an integrative taxonomy approach, we revealed and described a new species from Kyrgyzstan—Turanium losi Karpiński, Plewa & Hilszczański sp. nov. Distinguishing characters from closely related Turanium pilosum (Reitter, 1891) are presented and their ecological associations are discussed. The key characters, including the male terminalia, were examined by means of scanning electron microscopy. High-quality stacked photographs of the habitus of the specimens are presented for both species and their geographical distributions are mapped. While the new species shows stable morphological characters that allow its differentiation from T. pilosum and the COI genetic distance between them is approx. 3%, the different species delimitation methods gave discordant results. Although the new species remained unrecognized for so long, it seems that these cerambycids are common in the region and both can be considered potentially invasive as they are apparently highly polyphagous. It has also been documented that they occur sympatrically in Kyrgyzstan. Both the Bayesian and maximum likelihood analyses of COI sequences confirmed the monophyly of the genus Turanium with strong support (PP 1 and BS 90, respectively). Moreover, the recently revealed polyphyly of the tribe Callidiini was supported by our analyses and, consequently, the discussion on the establishment of a new tribe Ropalopini is raised. This study further corroborates the effectiveness of DNA barcoding as a tool in detecting new species and provides some of the first sequences for Central Asian cerambycids, which remain almost completely unknown in terms of molecular studies.

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DOI: 10.3897/asp.79.e64325

Authors: Massimo Meregalli, Manfred Kahlen, Riccardo Monguzzi, Valentina Marzia Rossi, Alfredo Santovito

Abstract: Dichotrachelus orobicus, a new species from the Bergamasque Prealps, is described. It is closely related to D. grignensis from the Grigna Massif, from which it differs mainly in the COI and ITS2 sequences, and minute morphological characters. Remarks on the possible epoch of speciation between the two taxa, with an analysis of the biogeographic scenario that may have led to the disjunction, are discussed.

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DOI: 10.3897/asp.79.e62554

Authors: Derek S. Sikes, Logan J. Mullen

Abstract: Abstract The omaliine rove beetle genus Phlaeopterus Motschulsky, 1853 contains 22 species. The genus is distributed across northwestern North America and eastern Asia. These beetles occur primarily along the edges of alpine snowfields and streams, habitats that are particularly sensitive to the impacts of climate change. Two species have not been collected since 1979 and 1984, one of which, Phlaeopterus bakerensis Mullen and Campbell, 2018, is a contender for the largest-bodied species among the over 1,600 species of the subfamily Omaliinae. Here, we present the first phylogeny of the genus, using Bayesian and maximum likelihood analyses based on DNA sequences from the mitochondrial gene COI and morphological data. We tested previous taxonomic hypotheses and most were rejected by all three analyses. Phlaeopterus castaneus Casey, 1893 is non-monophyletic based on COI sequences and may have hybridized with P. loganensis Hatch, 1957. We found support for the monophyly of the genus Phlaeopterus. Our analyses suggest the common ancestor of the genus had small-bodied adults (maximum body size under 5 mm) with ocelli. Within this small-bodied radiation of species, ocelli were lost once and there were two separate evolutionary transitions to large-bodied adults. Although all the large-bodied species are snowfield-associated and only 25% of the small-bodied species are, we did not find statistical support for a relationship between large body size and use of snowfield habitats. These findings represent the first modern phylogenetic reconstruction of species-level relationships within the rove beetle subfamily Omaliinae using both morphological and molecular data.

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DOI: 10.3897/asp.79.e66966

Authors: Stephanie Haddad, Nayeli Gutiérrez, Felipe A. Noguera, Seunggwan Shin, Petr Svacha, Duane D. McKenna

Abstract: Taxonomic placement of the enigmatic monotypic Mexican longhorned beetle genus Vesperoctenus Bates is examined through inclusion in and reanalysis of the dataset of Haddad et al. (2018, Systematic Entomology 43: 68–89). We describe and discuss the phylogenetic significance of the internal structures of a recently collected V. flohri female from the Sierra de la Laguna mountain range in Mexico, the same specimen from which phylogenomic data was generated. Our phylogenomic analyses (469 genes) recovered Vesperoctenus with maximal statistical support within the cerambyciform family Vesperidae, sister to Vesperus Dejean (Vesperinae). Vesperus + Vesperoctenus were recovered sister to Philinae, and collectively form a clade sister to Anoplodermatinae. Thus, we place V. flohri within Vesperidae: Vesperinae: Vesperoctenini based on analyses of large-scale phylogenomic data. Finally, we propose that the conservation status of V. flohri merits assessment.

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DOI: 10.3897/asp.79.e61499

Authors: Lech Karpiński, Patrick Gorring, Lech Kruszelnicki, Denis G. Kasatkin, Wojciech T. Szczepański

Abstract: This paper discusses ecological adaptation based on a case study of Anoplistes halodendri and Anoplistes kozlovi (Coleoptera: Cerambycidae) that occur in the arid zone of Mongolia. By applying an integrative taxonomy approach, we revealed one of the first documented cases of sympatrically occurring ecotypes in Polyphaga and the second case of ecotypes in the family Cerambycidae. The results of the analysis of the ecological data, molecular analysis of mitochondrial (COI) and nuclear (ArgK and CAD) genes, as well as a detailed morphological examination, which consisted of a study on the male genitalia including the endophallic structures, supported the hypothesis that these two entities, which are commonly considered separate species, represent only ecologically adapted forms that are associated with rocky hills and sandy/gravelly plains, respectively. Therefore, a synonym is restored: Anoplistes halodendri minutus Hammarström, 1892 = Asias kozlovi Semenov and Znojko, 1934, syn. res. The differences in the elytral pattern and shade appear to be adapted to the different substrates in these distinct habitats. A probable scenario assumes that these two forms arose in parapatry, independently in multiple populations, under parallel speciation during the intensification of the aridification across the region in the period during which the Gobi Desert was formed (~ 24 to 2.6 Ma) and they may evolve into separate species in the future. The phylogenetic relationships of some taxa representing the most closely related genera of the tribe Trachyderini were analysed and the questioned species status of Anoplistes jacobsoni was confirmed. Low interspecific differences in the endophallic structures in the genus Anoplistes and between some species of the genus Amarysius indicate that they are evolutionarily relatively young groups. The practical differences between ecotype and subspecies are also discussed.

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