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A new species of Bathynellidae is described from Colorado (USA). Hobbsinella gunnisonensis Camacho & Taylor sp. nov. displays a unique combination of morphological characters including seven-segmented antenna lacking medial seta on exopod, antennule slightly longer than antenna, three-segmented mandibular palp, four articles on endopod of thoracopods I to VII and five spines on sympod and three spines on endopod of the uropods. Partial sequences of cytochrome oxidase I (COI) and 18S have been obtained from several specimens of the new species. The mitocondrial and nuclear DNA data complement the traditional morphological taxonomic description support the validity of the new species. Molecular data for the Bathynellidae demonstrate the presence of two highly divergent genetic units, with the new species placed in the genus Hobbsinella. With the description of Hobbsinella gunnisonensis Camacho & Taylor sp. nov. and its molecular characterization, we discovered an interesting distribution of the genus, which occurs in both sides of the Continental Divide (Texas and Colorado) and different habitats.

A new species of Bathynellidae is described from Colorado (USA). Hobbsinella gunnisonensis Camacho & Taylor sp. nov. displays a unique combination of morphological characters including seven- segmented antenna lacking medial seta on exopod, antennule slightly longer than antenna, three-segmented mandibular palp, four articles on endopod of thoracopods I to VII and ve spines on sympod and three spines on endopod of the uropods. Partial sequences of cytochrome oxidase I (COI) and 18S have been obtained from several specimens of the new species. The mitocondrial and nuclear DNA data complement the traditional morphological taxonomic description support the validity of the new species. Molecular data for the Bathynellidae demonstrate the presence of two highly divergent genetic units, with the new species placed in the genus Hobbsinella. With the description of Hobbsinella gunnisonensis Camacho & Taylor sp. nov. and its molecular characterization, we discovered an interesting distribution of the genus, which occurs in both sides of the Continental Divide (Texas and Colorado) and different habitats.

Tok-tokkies are one of the most iconic lineages within Tenebrionidae. In addition to containing some of the largest darkling beetles, this tribe is recognized for its remarkable form of sexual communication known as substrate tapping. Nevertheless, the phylogenetic relationships within the group remain poorly understood. This study investigates the usefulness of female terminalia morphology for delimiting Sepidiini and reconstructing relationships among it. Data on the structure of the ovipositors, genital tubes and spicula ventrali have been generated for >200 species representing 28 Pimeliinae tribes. This dataset was used in a comparative analysis at the subfamilial level, which resulted in recognition of several unique features of tok-tokkie terminalia. Additionally, new features linking phenotypically challenging tribes also were recovered (Cryptochilini + Idisiini + Pimeliini).Secondly, 23 characters linked to the structure of female terminalia were defined for tok-tok beetles. Cladistic analysis demonstrates the nonmonophyletic nature of most of the recognized subtribes. The morphological dataset was analysed separately and in combination with available molecular data (CAD, Wg, cox1, cox2, 28S). All obtained topologies were largely congruent, supporting the following changes: Palpomodina Kaminski & Gearner subtr.n. is erected to accommodate the genera Namibomodes and Palpomodes; Argenticrinis and Bombocnodulus are transferred from Hypomelina to Molurina; 153 species and subspecies previously classified within Psammodes are distributed over three separate genera (Mariazofia Kaminski nom.n., Piesomera stat.r., Psammodes sens.n.). Psammodes sklodowskae Kaminski & Gearner sp.n. is described. Preliminary investigation of the ovipositor of Mariazofia basuto (Koch) comb.n. was carried out with the application of microcomputed tomography, illuminating the muscular system as a reliable reference point for recognizing homologous elements in highly modified ovipositors.

The article provides information on the organizational aspects of the International Tenebrionoidea Virtual Symposium VI and the preceding North American Tenebrionidae identification and classification virtual workshop. Both events took place in May of 2021. Useful references to permanent online resources (schedules, talks) from both these meetings are presented.

La familia Tenebrionidae (Latreille, 1802) representa uno de los grupos de coleópteros más diversos, con más de 30.000 especies descritas en todo el mundo, cuyo ancestro común se remonta al Jurásico. Para entender cómo se ha generado la diversidad actual de tenebriónidos, hemos seleccionado dos géneros con reducida capacidad dispersiva: Pimelia (Pimeliinae) y Misolampus (Stenochiinae). Nuestro estudio se centra en la Cuenca Mediterránea, donde ambos géneros concentran gran parte de sus endemismos y la distribución total de algunos de sus taxones. En esta tesis, se analizaron las relaciones filogenéticas dentro de ambos géneros basadas en marcadores mitocondriales y nucleares, discutiendo la taxonomía y sistemática de ambos grupos. Asimismo, se estableció un marco temporal de origen y diversificación, y se reconstruyeron los escenarios biogeográficos históricos más plausibles para profundizar en el conocimiento de su historia evolutiva y en los mecanismos responsables de su diversidad y distribución actual. A la luz de nuestros resultados, reevaluamos el estado de los subgéneros y especies de Pimelia y Misolampus, describiendo seis nuevos subgéneros e identificando nuevas especies crípticas. Además, situaron el antepasado de Pimelia en el norte de África como región más probable (Oligoceno Temprano). Interesados en cómo la compleja historia geológica de la Cuenca Mediterránea Occidental habría condicionado la diversificación de este grupo y la dispersión en las expansiones y contracciones de rango de sus especies, analizamos el papel del Estrecho de Gibraltar, una barrera biogeográfica histórica para muchos artrópodos. Pimelia y Misolampus, pese a su reducida capacidad dispersiva, tienen representantes en ambos lados del Estrecho. El estudio de los patrones de diversidad genética en Pimelia en el área Bético-Rifeña constató que, dentro de un mismo linaje, existían casos en los que el Estrecho no había representado una barrera biogeográfica, como en Amblyptera y Ambly

[Aim]: Discerning the relative role of geographical and ecological factors in promoting diversification is central to our understanding of the origin and maintenance of biodi-versity. We explore the roles of geology and ecological tolerance in the diversification of a group of wingless beetles with low dispersal potential. [Location]: Western Mediterranean (Iberian Peninsula and North Africa)., [Taxon]: Darkling beetles (Tenebrionidae: Misolampus)., [Methods]: We sequenced nine gene fragments from the mitochondrial and nuclear genomes in all extant Misolampus species to reconstruct their phylogeny, evaluate species boundaries and potential contact zones and estimate divergence times. We modelled species distributions for different time periods to infer ecological preferences and assess the effects of climatic changes since the last interglacial. We used a time-stratified process-based biogeographical model to estimate ancestral areas of origin and the evolution of geographical ranges., [Results]: The palaeoclimatic model projections show contractions of favourable areas during the last interglacial period and mid-Holocene, and wide stretches of suitable areas during the last glacial maximum. Analyses of ancestral bioclimatic preferences reveal ecological adaptations in isolated lineages within three species. The phylog-eny of Misolampus is strongly supported and unveils deep divergences within the six species. Two well-supported clades were recovered, one distributed in North Africa- Balearic Islands and another in the Iberian Peninsula. The divergence between the North African and Iberian clades occurred during the early Miocene. The geographical ranges of species are dynamic, shaped by biotic (e.g. life history, demography, species interactions) and abiotic (e.g. tectonic events, climatic variables) factors that vary through space and time. This variation ultimately dictates differential patterns of population viability, connectivity and growth through time in dif-ferent sections of species ranges, with important consequences for species diversification processes (Bellard et al., 2012; Gouveia et al., 2014; Thuiller et al., 2005; Wiens, 2011). Following climatic changes, species can respond by contracting their ranges into cli-matic refugia and/or dispersing to areas where their environmen-tal preferences are maintained (niche tracking), by adapting to new environmental conditions in situ (niche evolution), and often by undergoing extinction in environmentally unsuitable areas (Kozak & Wiens, 2006; Yesson & Culham, 2006a). The fragmentation of ancestral distributions because of tectonic or climatic changes pro-motes allopatric speciation, sometimes associated with niche evolution, whereas demographic expansion of populations tracking climatic changes can lead to range fusion and admixture between formerly isolated population groups. Discerning the relative role of biotic and abiotic factors in promoting diversification is central to our understa, [Main conclusions]: Our results favour a dual role of vicariance and dispersal in driving the historical biogeography and diversification of Misolampus since the early Miocene. We also found evidence for incipient speciation events, underscoring the role of tectonic events and adaptation to local climatic conditions in the diversification of the group.

The “Iberobathynella group”, or Iberobathynellini tribe, is a complex of six genera consisting of 33 nominal species and several cryptic species with an amphiatlantic distribution (in Europe, North Africa and North America). A modern systematic revision of this group of subterranean crustaceans is presented here. A phylogenetic and biogeographic study using morphological and molecular data (mitocondrial coi and nuclear 18S) was carried out and allowed to a) re-evaluate the taxonomic status and validity of previously erected subtribes, genera and subgenera that show congruence in the data; b) assess whether the identified mitochondrial lineages represent cryptic species; c) provide a plausible phylogenetic hypothesis for the relationships within Iberobathynellini and with the other two genera of the family Parabathynellidae that inhabit North America and Europe (Montanabathynella and Parabathynella, respectively); d) propose a plausible temporal and historical framework (paleobiogeographic scenario) for the diversification and evolution of the Iberobathynellini tribe based on the current distribution of morphotypes and their estimated times of divergence. Our results show that in parabathynellids, molecular and morphological divergences are not always congruent. Subtribe and subgenus are invalid categories so they must be eliminated. Paraiberobathynella genus needs to be revisited. The molecular dating results support the early divergence of the Iberobathynellini Tribe (Upper Cretaceous, around 78 Mya) and the vicariance by plate tectonics as main factor to explain the amphi-Atlantic distribution shown by this ancient subterranean crustacean group. Since there are species morphologically very similar to I. magna and I. imuniensis, but genetically different, we can ensure the existence of at least three cryptic species. Texanobathynella is undoubtedly a valid genus distinct from Iberobathynella. Montanabathynella and Parabathynella are two well-differentiated genera c

The subgenus Amblypteraca Mas-Peinado, Buckley, Ruiz & García-París, 2018 of Pimelia Fabricius, 1775, is restricted to the southern Iberian Peninsula and western Maghreb (northern and western Morocco). The distribution of Amblypteracathroughout the African-European edges overlaps largely with the geographic range of the subgenus Amblyptera,which is sister to the clade grouping subgenera Amblypteraca and Ecphoroma Solier, 1836. Delimiting species boundaries in the speciose genus Pimelia is often challenging, and the taxonomic status of some groups within the aforementioned subgenera is still a matter of debate. Here, we aim to stabilize some of the available names in Amblypteraca, and to correct some previous misidentifications. For that purpose, we discuss the composition and taxonomic structure within Amblypteraca by (i) assessing the phylogenetic congruence between mitochondrial and nuclear markers, and (ii) examining external morphological traits in 568 Amblypteraca specimens under the light of the phylogenetic hypotheses proposed here. Based on our results, Amblypteraca consists of three species: P. rotundipennis Kraatz, 1865, P. fairmairii Kraatz, 1865and P. chrysomeloides Pallas, 1781. Both molecular and morphological data revealed four lineages within P. chrysomeloides: P. chrysomeloideschrysomeloides, distributed on both sides of the Strait of Gibraltar; P. chrysomeloides fornicata Herbst, 1799 fromPortugal (Troia region); P. chrysomeloides bathyglypta Antoine, 1949, restricted to a narrow strip between Larache and Arbaoua (northern atlantic Moroccan coast),and P. chrysomeloides subris Koch, 1941 from Kenitra-La Mamora forest (Morocco). We designate a neotype of Tenebrio chrysomeloides Pallas, 1781 and propose the synonymy of P. chrysomeloides (Pallas, 1781) = P. obesa Solier, 1836 syn. n. Pimelia tristis Haag-Rutenberg, 1875,previously misidentified andincluded in Amblypteraca, is now transferred back to Amblyptera. Further studies with ad hoc sampling desi

The subgenus Amblypteraca Mas-Peinado, Buckley, Ruiz & García-París, 2018 ofPimelia Fabricius, 1775, is restricted to the southern Iberian Peninsula and western Maghreb (northern and western Morocco). The distribution of Amblypteraca throughout the African-European edges overlaps largely with the geographic range of the subgenus Amblyptera, which is sister to the clade grouping subgenera Amblypteraca and Ecphoroma Solier, 1836. Delimiting species boundaries in the speciose genusPimelia is often challenging, and the taxonomic status of some groups within the aforementioned subgenera is still a matter of debate. Here, we aim to stabilize some of the available names in Amblypteraca, and to correct some previous misidentifications. For that purpose, we discuss the composition and taxonomic structure within Amblypteraca by (i) assessing the phylogenetic congruence between mitochondrial and nuclear markers, and (ii) examining external morphological traits in 568 Amblypteraca specimens under the light of the phylogenetic hypotheses proposed here. Based on our results, Amblypteraca consists of three species: P. rotundipennis Kraatz, 1865, P. fairmairii Kraatz, 1865 and P. chrysomeloides Pallas, 1781. Both molecular and morphological data revealed four lineages within P. chrysomeloides: P. chrysomeloides chrysomeloides, distributed on both sides of the Strait of Gibraltar; P. chrysomeloides fornicata Herbst, 1799 from Portugal (Troia region); P. chrysomeloides bathyglypta Antoine, 1949, restricted to a narrow strip between Larache and Arbaoua (northern atlantic Moroccan coast), and P. chrysomeloides subris Koch, 1941 from Kenitra-La Mamora forest (Morocco). We designate a neotype of Tenebrio chrysomeloides Pallas, 1781 and propose the synonymy of P. chrysomeloides (Pallas, 1781) = P. obesa Solier, 1836 syn. n. Pimelia tristis Haag-Rutenberg, 1875, previously misidentified and included in Amblypteraca, is now transferred back toAmblyptera. Further studies with ad hoc sampling

Three new species of Parabathynellidae (Crustacea: Bathynellacea) are described using morphological and molecular data, and addi-tional populations identified as Texanobathynella cf. bowmani are placed within a molecular phylogeny for the group. The finding of Montanabathynella pecosensis sp. nov. entails a southern range expansion of over 1500 km for the genus, previously thought to be restricted to Montana, USA. Montanabathynella pecosensis sp. nov. displays a unique combination of morphological features, including eight main teeth on the labrum; four and six teeth on the pars incisiva and pars molaris of the mandible, respectively; eight teeth on the distal endite of MxI; epipod present on ThII and III; four teeth on the first segment of female ThVIII; and an unpro-nounced anal operculum. The other two new species described here belong to the genus Texanobathynella Delamare Deboutteville et al., 1975, previously consisting of only two species. Both species, Texanobathynella aaronswinki sp. nov and T. coloradoensis sp. nov., display combinations of morphological features distinct from the other two previously known species of the genus. Mitochondrial (COI) and nuclear (18S) markers support the validity of the new species and enable us to place them, along with T. cf. bowmani, within a larger phylogenetic framework of the Parabathynellidae.

Bathynellidae is the neglected family of Bathynellacea, a groundwater group of crus- taceans with 33 genera and 107 species described and almost half of them included in the `catch-all¿ genus Bathynella. Due to the morphological homogeneity of the spe- cies, the taxonomic uncertainties have accumulated over time. Therefore, to explore the phylogenetic relationships among taxa, a combined approach using morphologi- cal and molecular data is needed. In this paper, we performed a phylogenetic analysis based on partial sequences of COI and 18S including 30 species of Bathynellidae. This data set represents the most updated one, including a new genus and a new spe- cies (Altainella calcarata gen. n. sp. n.) of Bathynellidae described here from Altai Mountain (Russia). The new taxon presents a new type of sexual dimorphisms with a peculiar structure on the coxopod of the male thoracopod VII and also sexual dimor- phism on the mandibular palp, as some species of the subfamily Gallobathynellinae. This is the first species of Bathynellidae that presents both variants of sexual dimor- phism. The molecular phylogeny supports the morphology and the erection of the new genus and displays highly divergent genetic units corresponding to subfamilies, with the new genus closer to Bathynellinae genera than other subfamilies. With the description of this new species and its molecular characterization, the existence of the other genera beyond Bathynella is confirmed in Eurasia.

Hunchback darkling beetles of the Ibero-Maghrebian genus Misolampus Latreille, 1807 (Tenebrionidae, Stenochiinae) encompass six species: M. gibbulus (Herbst 1799), M. goudotii Guérin-Méneville, 1834, M. lusitanicus Brême, 1842, M. ramburii Brême, 1842, M. scabricollis Graells, 1849, and M. subglaber Rosenhauer, 1856. Previously known distribution ranges of the species were delineated using many old records, the persistence of such populations being questionable under the current situation of global biodiversity loss. Additionally, the status of geographically isolated populations of the genus have been the subject of taxonomic controversy. An exhaustive bibliographical revision and field search was undertaken, and the Misolampus collection of the Museo Nacional de Ciencias Naturales (MNCN-CSIC) was revised. The aims are to (i) provide an updated geographic distribution range for the species of Misolampus; (ii) to determine the taxonomic status of controversial populations; (iii) to provide a catalogue for Misolampus; and (iv) to discuss the conservation status of these saproxylic beetles. As a result, a catalogue including synonymies and type localities, geographical records, diagnoses, and information on natural history for all species of Misolampus is presented. The results reveal that the distribution ranges of the species of Misolampus have not undergone a reduction in the last century, and indicate the presence of the genus in areas where it had never been recorded before. The morphological variability of M. goudotii drove the proposal of different taxa that are here formally synonymised as follows: M. goudotii Guérin-Méneville, 1834 = M. erichsoni Vauloger de Beaupré, 1900, syn. nov. = M. peyerimhoffi Antoine, 1926, syn. nov.

Forficula auricularia (the European earwig) is possibly a complex of cryptic species. To test this hypothesis, we performed: (1) a phylogeographic study based on fragments of the mitochondrial COI and the nuclear ITS2 markers on a wide geographic sampling, (2) morphometric analyses of lineages present in Spain and (3) niche overlap analyses. We recovered five reciprocally monophyletic ancient phylogroups with unique historical patterns of distribution, climatic niches and diversification. External morphology was conserved and not correlated with speciation events, except in one case. Phylogenetic placement of the morphologically distinct taxon renders F. auricularia paraphyletic. Based on the congruence of the phylogenetic units defined by mtDNA and nuclear sequence data, we conclude that phylogroups have their own historical and future evolutionary trajectory and represent independent taxonomic units. Forficula auricularia is a complex of at least four species: the morphologically diagnosable Forficula aeolicaGonzález-Miguéns & García-París sp. nov., and the cryptic taxa: Forficula mediterranea González-Miguéns & García-París sp. nov., Forficula dentataFabricius, 1775 stat. nov. and Forficula auriculariaLinnaeus, 1758 s.s. New synonymies of F. dentata are also proposed.

A preliminary molecular phylogenetic framework for 12 genera (23 species) of the family Parabathynellidae from Europe, Australia, North Africa and India is presented based on mitochondrial and nuclear markers (Cox1 and 18S). The generated hypothesis places the Southeast Asia genus Paraeobathynella closer to European genera (Iberobathynella, Paraiberobathynella and Parabathynella) than to the Australian (Brevisomabathynella, Atopobathynella, Billibathynella, Octobathynella, Arkaroolabathynella and Lockyerenella) or Indian genera (Habrobathynella), or to the cosmopolitan genus Hexabathynella (Spain and Australia). Three new species of the genus Paraeobathynella from Thailand, P. ratensis n. sp., P. siamensis n. sp. and P. hanjavanitiana n. sp., are described based on morphological and molecular features. This is the first record of the genus from Thailand and extends its range of distribution within Asia, where it was previously known only from Vietnam. The new species are clearly separated as independent units at least since the Middle Miocene.

The genus Pimelia Fabricius is the most speciose in the tribe Pimeliini (Coleoptera: Tenebrionidae). It consists of approximately 320 species distributed mainly in xeric environments in the western Palaearctic region and northern deserts in the Afrotropical region. Pimelia is a very heterogenous group despite its conserved morphology. Based on two mitochondrial (cox1 and 16S) and five nuclear markers (ITS1, ITS2, 18S, Wg and H3), we carried out a phylogenetic and biogeographic study to: (i) provide a comprehensive phylogenetic hypothesis for the relationships within Pimelia; (ii) re‐evaluate the taxonomic status and organization of clades (subgenera) within the genus; and (iii) establish a temporal framework and historical scenario for the evolution and diversification of Pimelia in the western Mediterranean Region. Pimelia presents a high degree of morphological conservatism and possibly stasis, often resulting in complexes of cryptic species. Repetitive patterns of variation are found recurrently between and within groups, which resulted in a conundrum of nonmonophyletic taxonomic assemblages based on morphological homoplastic characters. Based on our results, we propose a new taxonomic arrangement for Pimelia with 14 clades, treated as subgenera, with the description of six new subgenera: Iberomelia subgen.n., Italomelia subgen.n., Magrebmelia subgen.n., Hispanomelia subgen.n., Massadraamelia subgen.n. and Amblypteraca subgen.n. Our results also indicate an Oligocene (Rupelian) origin for Pimelia, and North Africa as the most likely area of origin. The complex geological and climatic history in the western Mediterranean region has shaped the distribution and diversity of species in the area. The current faunal assemblages observed in Pimelia are the result of complex patterns of recurrent processes of dispersal and vicariance, including in situ diversification through time in different areas of the Mediterranean basin. Pimelia is a fascinating yet very challengin

Bathynella natans Vejdovsky, 1882, the first described Bathynellidae species, was found in a well in Prague. Its original description was short and incomplete, and the holotype was already damaged in 1899. The lack of type material resulted in the poor description of the type genus of the morphologically very homogeneous family. This troubled beginning is the basis of a taxonomic problem that has led to the assignment of many species in Europe and other continents to the genus Bathynella. As currently understood, the genus is cosmopolitan, with more than 50 species and subspecies, most of them insufficiently described. We analyse the systematic position of the 29 genera and approximately 100 species. This analysis includes genera from Europe, USA and Western Australia, which have been characterized with molecular data (18S and COXI). In most cases, the genetic distances among genera and species are congruent with detected morphological differences. This is the first world molecular phylogenetic reconstruction of Bathynellidae, and it creates a framework for (1) confirming or discarding the cosmopolitan status of the genus Bathynella, (2) clarifying taxonomic uncertainties, (3) adequately describing new species, (4) advancing knowledge of the family’s real diversity and (5) describing the phylogenetic relationships among genera from different continents.

Intra-specific coloration polymorphism coupled with an ancient process of lineage differentiation in Berberomeloe majalis (Linnaeus, 1758) offers the opportunity to analyse the temporal scenario in which the correlation between toxicity and coloration might have evolved. Based on phylogenetic and phylogeographic analyses, we identified the timing for the split between red-striped and entirely black morphotypes of B. majalis. To evaluate whether coloration patterns and toxicity are related in this species, we quantified the concentration of cantharidin across morphotypes and phylogeographic lineages. Phylogenetic analyses based on cox1 mitochondrial DNA sequences recovered three major clades where both morphotypes were intermingled, indicating a multiple homoplastic origin for the entirely black coloration. Our analyses showed that cantharidin content did not differ between morphs of B. majalis; however, it significantly increased in haemolymph in females kept isolated from males, which reveals the females’ ability either to concentrate cantharidin towards haemolymph or to synthesize cantharidin themselves. Lack of monophyly and absence of genetic isolation in both morphotypes favour the hypothesis of a recent homoplastic phenomenon to explain the loss of the striped pattern. Our phylogenetic and phylogeographic analyses show that changes in coloration are recent, suggesting that the ancient pressures that fixed and maintained red-striped colorations are no longer acting today on B. majalis. The absence of change in cantharidin content (i.e. entirely black and red-striped specimens are equally poisonous) suggests that the evolution of colour polymorphisms in B. majalis is probably decoupled from toxicity.