On the presence of Scaurus puncticollis Solier, 1838 (Coleoptera: Tenebrionidae) in Qatar

As part of the biodiversity surveys conducted by the Department of Biodiversity at the Qatar Environment and Energy Research Institute (QEERI), we can report the presence of Scaurus puncticollis Solier, 1838 in Qatar, and confirm the first record for the tribe Scaurini Billberg, 1820 in the country. We found two living specimens and remains of a third one distributed over the year (February, April and October) in three different locations of the Al Rayyan Municipality in Qatar. These findings reveal that the fauna of darkling beetles of Qatar is, despite previous efforts, still underestimated. To document and report the actual Biodiversity Heritage of Qatar, new exhaustive field biodiversity surveys should be conducted to complete the catalogue of Tenebrionoidae for the State of Qatar.Peer Reviewe

Geology-based and ecological processes of divergence between and within species of wingless darkling beetles

Aim: Discerning the relative role of geographical and ecological factors in promoting diversification is central to our understanding of the origin and maintenance of biodiversity. 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 phylogeny 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. Biogeographical analyses infer an ancestral range including the Iberian, Betic and Rifean Plates, with subsequent splits followed by dispersal events. 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.Visit of PMP to the Institut des Sciences de l'Evolution de Montpellier (ISEM) was supported by the Ministerio de Ciencia, Innovación y Universidades, Spain (MICINN-FEDER) through contract BES-2016-077777. This study was supported by the Spanish government (MINECO/MIUC/AEI) and the European Fund for Regional Development (FEDER) under grants CGL2015-66571-P (collecting and Museum visits) and PID2019-110243GB-I00/AEI/10.13039/501100011033 (Ministerio de Ciencia, Innovación y Universidades, Spain) (molecular analyses) to MGP. Support of the publication fee was granted by the CSIC Open Access Publication Support Initiative through its Unit of Information Resources for Research (URICI)

A preliminary report on the distribution of lizards in Qatar

We have updated the list of the lizard species present in Qatar and produced the first distribution maps based on two field surveys in 2012 and 2013. We used the QND95/Qatar National Grid with a grid of 10 × 10 km squares for mapping. Our results show the occurrence of 21 lizard species in Qatar, from the 15 species indicated in the last biodiversity report conducted in 2004. The most abundant family found in Qatar is Gekkonidae with nine species (Bunopus tuberculatus, Cyrtopodion scabrum, Hemidactylus robustus, H. flaviviridis, H. persicus, Stenodactylus arabicus, S. slevini, S. doriae, Pseudoceramodactylus khobarensis), followed by Lacertidae with four species (Acanthodactylus schmidti, A. opheodurus, Mesalina brevirostris, M. adramitana), Agamidae with three species (Trapelus flavimaculatus, Uromastyx aegyptia, Phrynocephalus arabicus), Scincidae with two species (Scincus mitranus, Trachylepis septemtaeniata), and Varanidae (Varanus griseus), Sphaerodactylidae (Pristurus rupestris) and Trogonophiidae (Diplometopon zarudnyi) with one species each. The species richness fluctuated largely across Qatar between one and eleven species per grid square. We believe that the lizard fauna records in Qatar are still incomplete and that additional studies are required. However, our study here fills a gap concerning lizard biodiversity knowledge in the Gulf Region. © Dan Cogǎlniceanu et al.Peer Reviewe

Female terminalia morphology and cladistic relations among Tok-Tok beetles (Tenebrionidae: Sepidiini)

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.Funding was provided by the National Science Centre, Poland (OPUS-19 Project 2020/37/B/NZ8/02496) and National Science Foundation (DEB-1754630/2009247)Peer reviewe

Evolutionary processes and speciation patterns in the Mediterranean Basin darkling beetles

Tesis doctoral inédita leída en la Universidad Autónoma de Madrid, Facultad de Ciencias, Departamento de Biología. 342 pp.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 Amblypteraca. Ambos subgéneros comparten patrones de colonizaciones post-Messinienses recurrentes a través del Estrecho, posiblemente impulsadas por eventos estocásticos o catastróficos. En otros subgéneros, como en Magrebmelia, el Estrecho habría representado una fuerte barrera al flujo génico desde el Mioceno Medio. Tras actualizar en un estudio el rango de distribución de las especies de Misolampus, se confirmó que el área límite entre África y Europa ha constituido una barrera para el intercambio génico desde el Mioceno temprano, sin existir evidencias de dispersión transcontinental desde entonces. Nuestros resultados situaron al ancestro común entre las placas Ibérica y BéticaRifeña como región más probable. Las especies de Misolampus han permanecido como linajes relictos desde el Mioceno, con expansiones y contracciones de rango durante el Pleistoceno que han dejado huellas de procesos de especiación incipientes. Tanto para Misolampus como para Pimelia el área Ibero-Magrebí ha servido como refugio y como centro de diversificación. Los resultados obtenidos en esta tesis constatan como la compleja historia geológica de la Cuenca Mediterránea Occidental desde el Oligoceno ha condicionado la diversificación y actual diversidad de Tenebrionidae en el área. Además, nuestros estudios filogenéticos y escenarios biogeográficos plantean también un escenario de partida para futuros estudios en ambos géneros, que pueden ayudarnos a comprender su resiliencia durante millones de años.Peer reviewe

Evolutionary processes and speciation patterns in the Mediterranean Basin darkling beetles

Tesis doctoral inédita leída en la Universidad Autónoma de Madrid, Facultad de Ciencias, Departamento de Biología. Fecha de lectura: 14-01-2022Esta tesis tiene embargado el acceso al texto completo hasta el 14-07-202

Cantharidin is conserved across phylogeographic lineages and present in both morphs of Iberian Berberomeloe blister beetles (Coleoptera, Meloidae)

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.CGL2010-15786, CGL2008-02567, CGL2012-36345, y CGL2015-66571-PPeer reviewe

The Strait of Gibraltar is an ineffective palaeogeographic barrier for some flightless darkling beetles (Coleoptera: Tenebrionidae: Pimelia)

The geographic distribution of a species is shaped by its biology and by environmental and palaeogeographic factors that interact at different spatial-temporal scales, which leads to distributions and diversification patterns observed between and within lineages. The darkling beetle genus Pimelia has been diversifying for more than 31.2 Mya showing different colonization patterns after the opening of the Gibraltar Strait 5 Mya. Three of the 14 subgenera of Pimelia have populations on both sides of the Strait. Through extensive sampling and the analysis of three molecular markers, we determine levels of intra- and interspecific genetic variation, identify evolutionary lineages in subgenera, estimate their temporal origin and distribution ranges and discuss the historical basis for the geographic and diversification patterns of Pimelia around the Strait. This single geographical feature acted both as a barrier and as a dispersal route for different Pimelia species. The Strait has represented a strong barrier for the subgenus Magrebmelia since the Middle Miocene. However, the subgenera Amblyptera and Amblypteraca share repetitive signatures of post-Messinian colonization across the Strait, possibly driven by stochastic or ‘catastrophic’ events such as tsunamis. Our demographic analyses support Wallace’s hypothesis on insect dispersal stochasticity. Some taxonomic changes, including the designation of a lectotype for Pimelia maura, are also proposed.Peer reviewe

Recurrent diversification patterns and taxonomic complexity in morphologically conservative ancient lineages of Pimelia (Coleoptera: Tenebrionidae)

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 challenging group, and the phylogenetic and systematic framework developed here will pave the way for further ecological, taxonomic, and evolutionary studies at local and regional scales.PMP is supported by MINECO‐FEDER through contract BES‐2016‐077777. This work has been funded by the project‐grant CGL2015‐66571‐P (MINECO‐FEDER) to MGP.Peer Reviewe

Taxonomy of the North Moroccan and Iberian species of the subgenusAmblypteraca (Coleoptera: Tenebrionidae: Pimeliinae: Pimelia)

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 designs and analytical tools would be in need to delimit the exact geographic ranges of these taxa, and to analyse the patterns of diversity within and among species and subspecies.Visit of PMP to the Magyar Természettudományi Múzeum of Budapest (HU-TAF-4091) and MGP to the Muséum National d’Histoire Naturelle was supported by the European Project SYNTHESYS (http://www.synthesys.info/). PMP is supported by MICINN-FEDER through contract BES-2016-077777 (Ministerio de Ciencia, Innovación y Universidades, Spain). This study was supported by the Spanish government (MINECO/MIUC/AEI) and the European Fund for Regional Development (FEDER) under grants Ministerio de Economía y Competitividad under Grant (collecting and Museum visits) and PID2019-110243GB-I00 / AEI/10.13039/501100011033 (Ministerio de Ciencia, Innovación y Universidades, Spain) (molecular analyses) to MGP.Peer reviewe