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The previously known spider beetle fauna of Socotra comprises four endemic species (Mezium erinaceus Belles, Sphaericus hirsutus Belles, Ptinus bertranpetiti Belles, and Silisoptinus inermicollis Belles) and Dignomus mesopotamicus (Pic), which is also known from Arabia and Mesopotamia. The study of new material has resulted in the discovery of a new genus and two new species, and many new localities of the previously known species. Here, a synopsis is presented that brings together all information available on the fauna of spider beetles of the Socotra archipelago, including updated distributions and ecological and biogeographical considerations of all species. http://www.zoobank.org/urn:lsid:zoobank.org:pub:3EB99992-B352-42AD-A33D-C01EF494666C. ; This work was supported by the Catalan Government (grants 2014 SGR 619, and 2017 SGR 1030).

[EN] Joaquim Mateu Sanpere was born in Barcelona on January 9, 1921 within a wealthy family of the Catalan bourgeoisie of the early twentieth century. His parents were Cristòfol Mateu Ferrer and Teresa Sanpere Cantelis, and the family had an illustrious ancestor, the historian, critic and politician Salvador Sanpere i Miquel, the father of Teresa Sanpere who had died in 1915. Joaquim was the second of the couple ́s four children: Eduard had preceded him, while Josep and Elena were born after him. ; [ES] Joaquím Mateu Sanpere nació en Barcelona el 9 de enero de 1921 en el seno de una familia acomodada de la burguesía catalana de principios del siglo XX. Sus padres eran Cristòfol Mateu Ferrer y Teresa San - pere Cantelis, y la familia contaba con un antepasado ilustre, el historiador, crítico y político Salvador Sanpere i Miquel, que fue el padre de Teresa Sanpere y murió en 1915. Joaquín fue el segundo de los cuatro hijos que tuvo la pareja. Eduard le había precedido, mientras que Josep y Elena nacieron después de él. ; [CA] Joaquim Mateu Sanpere va néixer a Barcelona el 14 de gener de 1921 en el si d'una família benestant de la burgesia catalana del principi del segle XX. Els seus pares eren Cristòfol Mateu Ferrer i Teresa Sanpere Cantelis, i la família tenia un avantpassat il·lustre, l'historiador, crític i polític Salvador Sanpere i Miquel, que va ser el pare de Teresa Sanpere i va morir el 1915. Joaquim va ser el segon dels quatre fills que va tenir la parella. Eduard l'havia precedit, mentre que Josep i Elena van néixer després d'ell. ; [FR] Joaquim Mateu Sanpere naquit le 9 janvier 1921 au sein d'une famille de la bourgeoisie catalane du début du XX e siècle. Ses parents étaient Cristòfol Mateu Ferrer et Teresa Sanpere Cantelis. La famille avait un ancêtre illustre en la personne de l'historien, critique et homme politique Salvador Sanpere i Mi - quel, qui était le père de Teresa Sanpere et mourut en 1915. Des quatre enfants du couple, Joaquim fut le deuxième, l'aîné étant Eduard et les plus jeunes, Josep et Elena. ; Peer reviewed

Wings were a fundamental morphological innovation for the adaptive radiation of insects, the most diversified group among all animals. Pterygote insects have two pairs of wings, the mesothoracic (T2) forewings and the metathoracic (T3) hindwings, whereas the prothorax (T1) is wingless. Using RNA interference approaches, we have found that the gene Sex combs reduced (Scr) determines the wingless identity of T1 in the cockroach Blattella germanica. Interference of Scr triggers the formation of ectopic wing structures in T1, which are formed from the expansion of the latero-posterior region of the pronotum, along with a contribution of the epimeron, a pleurite of T1. These data support the theory of a dual origin for insect wings, from pronotal (tergal origin theory) and pleural (pleural origin theory) structures and genes. ; Support for this research was provided to X.B. by the Spanish Ministry of Science and Innovation (grant no. CGL2008-03517/BOS) and Ministry of Economy and Competitiveness (grant CGL2012-36251 and CGL2015-64727- P, including FEDER funds) and by the Catalan Government, AGAUR (2014 SGR 619). M.E.N. was awarded a post-doctoral fellowship by the Fundação de Amparo à Pesquisa do Estado de São Paulo, Brazil (FAPESP 2012/15397-0). ; Peer reviewed

Juvenile Hormone (JH) represses metamorphosis of young instars in insects. One of the main players in hormonal signalling is Methoprene-tolerant (Met), which plays the role of JH receptor. Using the Polyneopteran insect Blattella germanica as the model and RNAi for transcript depletion, we have confirmed that Met transduces the antimetamorphic signal of JH in young nymphs and plays a role in the last nymphal instar moult in this species. Previously, the function of Met as the JH receptor had been demonstrated in the Eumetabola clade, with experiments in Holometabola (in the beetle Tribolium castaneum) and in their sister group Paraneoptera (in the bug Pyrrhocoris apterus). Our result shows that the function of Met as JH receptor is also conserved in the more basal Polyneoptera. The function of Met as JH transducer might thus predate the evolutionary innovation of metamorphosis. Moreover, expression of Met was also found in last nymphal instar of B. germanica, when JH is absent. Depletion of Met in this stage provoked deficiencies in wing growth and ecdysis problems in the imaginal moult. Down-regulation of the ecdysone-inducible gene E75A and Insulin-Like-Peptide 1 in these Met-depleted specimens suggest that Met is involved in the ecdysone and insulin signalling pathways in last nymphal instar, when JH is virtually absent. © 2014 Lozano, Belles. ; Financial support for this research was provided by the Spanish MICINN (grant CGL2008-03517/BOS to XB and predoctoral fellowship to JL) and MINECO (grant CGL2012-36251 to XB), and from the Catalan Government (2009 SGR 1498). The research has also benefited from FEDER funds. ; Peer Reviewed

The success of neopteran insects, with 1 million species described, is associated with developmental innovations such as holometaboly and the evolution from short to long germband embryogenesis. To unveil the mechanisms underlining these innovations, we compared gene expression during the ontogeny of two extreme neopterans, the cockroach Blattella germanica (polyneopteran, hemimetabolan, and short germband species) and the fly Drosophila melanogaster (endopterygote, holometabolan, and long germband species). Results revealed that genes associated with metamorphosis are predominantly expressed in late nymphal stages in B. germanica and in the early-mid embryo in D. melanogaster. In B. germanica the maternal to zygotic transition (MZT) concentrates early in embryogenesis, when juvenile hormone factors are significantly expressed. In D. melanogaster, the MZT extends throughout embryogenesis, during which time juvenile hormone factors appear to be unimportant. These differences possibly reflect broad trends in the evolution of development within neopterans, related to the germband type and the metamorphosis mode. ; This work was supported by the Spanish Ministry of Economy and Competitiveness (grants CGL2012-36251 and CGL2015-64727-P to X.B., including FEDER funds), Spanish Ministry of Science and Innovation (grants BFU2011–22404 and CGL2016–76011-R to M.-D.P., including FEDER funds), and Catalan Government (grants 2014 SGR 619 and 2017 SGR 1030). ; Peer reviewed

[Background]: Do miRNAs contribute to specify the germ-band type and the body structure in the insect embryo? Our goal was to address that issue by studying the changes in miRNA expression along the ontogeny of the German cockroach Blattella germanica, which is a short germ-band and hemimetabolan species. [Results]: We sequenced small RNA libraries representing 11 developmental stages of B. germanica ontogeny (with especial emphasis on embryogenesis) and the changes in miRNA expression were examined. Data were compared with equivalent data for two long germ-band holometabolan species Drosophila melanogaster and Drosophila virilis, and the short germ-band holometabolan species Tribolium castaneum. The identification of B. germanica embryo small RNA sequences unveiled miRNAs not detected in previous studies, such as those of the MIR-309 family and 54 novel miRNAs. Four main waves of miRNA expression were recognized (with most miRNA changes occurring during the embryonic stages): the first from day 0 to day 1 of embryogenesis, the second during mid-embryogenesis (days 0-6), the third (with an acute expression peak) on day 2 of embryonic development, and the fourth during post-embryonic development. The second wave defined the boundaries of maternal-to-zygotic transition, with maternal mRNAs being cleared, presumably by Mir-309 and associated scavenger miRNAs. [Conclusion]: miRNAs follow well-defined patterns of expression over hemimetabolan ontogeny, patterns that are more diverse during embryonic development than during the nymphal stages. The results suggest that miRNAs play important roles in the developmental transitions between the embryonic stages of development (starting with maternal loading), during which they might influence the germ-band type and metamorphosis mode. ; This work was supported by the Spanish Ministry of Economy and Competitiveness (grants CGL2012–36251 and CGL2015–64727-P to X.B.), the Spanish Ministry of Science and Innovation (grants BFU2011–22404 and CGL2016–76011-R to M.D.P.) and the Catalan Government (2014 SGR 619). It also received financial assistance from the European Fund for Economic and Regional Development (FEDER funds to X.B. and M.D.P.). ; We acknowledge support of the publication fee by the CSIC Open Access Publication Support Initiative through its Unit of Information Resources for Research (URICI). ; Peer Reviewed

Orcokinins (OKs) are neuropeptides that were first identified in crustacean through their myotropic activity. In insects, the OK gene gives rise to two mRNAs coding for two different families of conserved mature neuropeptides: OKA and OKB. Although OKs are conserved in many insect species, its physiological role in this animal class is not fully understood. Until now prothoracicotropic, regulatory of light entrainment to the circadian clock and >awakening> activities have been reported for these peptides in different insect species. Here we report the identification of OKA and OKB precursors in the cockroach Blattella germanica. OKA mRNA was detected in brain, whereas OKB mRNA was detected both in brain and midgut. In vivo silencing of OK precursors suggests the involvement of OK gene products in the regulation of vitellogenin expression in the fat body, an action that appears to be independent of juvenile hormone. This is the first time that a function of this kind has been reported for OKs. ; This work was supported by the Spanish MICINN (Grants CGL2008-03517/BOS to XB and BFU2010-15906 to JLM) and MINECO (Grant CGL2012-36251 to XB), and Catalan Government (2014 SGR 619). The research has also benefited from FEDER funds. SO was recipient of an external fellowship from National Council of Scientific and Technical Research (CONICET-Argentina) to work in the Institute of Evolutionary Biology, in Barcelona. ; Peer Reviewed

Current approaches for insect gene editing require microinjection of materials into early embryos. This severely limits the application of gene editing to a great number of insect species, especially to those whose reproduction systems preclude access to early embryos for injection. To overcome these limitations, we report a simple and accessible method for insect gene editing, termed "direct parental" CRISPR (DIPA-CRISPR). We show that injection of Cas9 ribonucleoproteins (RNPs) into the haemocoel of adult females efficiently introduces heritable mutations in developing oocytes. Importantly, commercially available standard Cas9 protein can be directly used for DIPA-CRISPR, which makes this approach highly practical and feasible. DIPA-CRISPR enables highly efficient gene editing in the cockroaches, on which conventional approaches cannot be applied, and in the model beetle Tribolium castaneum. Due to its simplicity and accessibility, DIPA-CRISPR will greatly extend the application of gene editing technology to a wide variety of insects. ; This research was supported by JSPS KAKENHI (nos. 20H02999 and 20K21311 to T.D.); JSPS Open Partnership Joint Research Projects (no. JPJSBP120209917 to T.D.); Spanish Ministry of Innovation and Competitiveness (PID2019-104483GB-I00 to X.B.); CSIC-Spain (2019AEP028 to M.-D.P.); and in part by Cabinet Office, Government of Japan, Cross-ministerial Moonshot Agriculture, Forestry and Fisheries Research and Development Program (no. JPJ009237 to T.D.). Y.S. is a JSPS Research Fellow with a research grant (DC1, 21J20658). ; Peer reviewed

Is there a correlation between miRNA diversity and levels of organismic complexity? Exhibiting extraordinary levels of morphological and developmental complexity, insects are the most diverse animal class on earth. Their evolutionary success was in particular shaped by the innovation of holometabolan metamorphosis in endopterygotes. Previously, miRNA evolution had been linked to morphological complexity, but astonishing variation in the currently available miRNA complements of insects made this link unclear. To address this issue, we sequenced the miRNA complement of the hemimetabolan Blattella germanica and reannotated that of two other hemimetabolan species, Locusta migratoria and Acyrthosiphon pisum, and of four holometabolan species, Apis mellifera, Tribolium castaneum, Bombyx mori and Drosophila melanogaster. Our analyses show that the variation of insect miRNAs is an artefact mainly resulting from poor sampling and inaccurate miRNA annotation, and that insects share a conserved microRNA toolkit of 65 families exhibiting very low variation. For example, the evolutionary shift toward a complete metamorphosis was accompanied only by the acquisition of three and the loss of one miRNA families. ; This work was supported by the Spanish Ministry of Economy and Competitiveness (grants CGL2012-36251 and CGL2015-64727-P to X.B.), Spanish Ministry of Science and Innovation (grant BFU2011-22404 to M.D.P.) and Catalan Government (2014 SGR 619). The research has also benefited from European Fund for Economic and Regional Development (FEDER) funds. B.F. was supported by the South-Eastern Norway Regional Health Authority grant #2014041. ; Peer reviewed

In adult females of the cockroach Blattella germanica, juvenile hormone (JH) promotes vitellogenin production. Depletion of Methoprene-tolerant (Met) mRNA levels with RNAi resulted in a clear reduction of vitellogenin expression in the fat body while basal oocyte growth was impaired. This demonstrates that Met is essential to transduce the vitellogenic signal of JH in this species. Interestingly, the expression of the transcription factor Kruppel homolog 1 (Kr-h1) was also reduced in Met-depleted specimens. This indicates that the JH signaling pathway promotes the transcription of Kr-h1 in adult fat body tissues, as occurs in the epidermis during nymphal development. Treatments with JH show that the expression of Met does not depend on JH, which suggests that Met is upstream the JH signaling pathway acting as JH receptor, as reported in other models and processes, especially in metamorphosis. JH treatments increased the transcription levels of vitellogenin and Kr-h1, which again suggests that Kr-h1 is a JH-dependent transcription factor in the fat body of adult females. The important roles of Met in nymphal development, as previously reported, and those reported herein in relation to reproduction, suggests that it can be an interesting target for the control of B. germanica in urban environments, using RNAi approaches. ; This work was supported by the Spanish Ministerio de Economía y Competitividad (grants CGL2012-36251 and CGL2015-64727-P to X.B., and BFU2010-15906 to JLM), and Catalan Government (2014 SGR 619). The research has also benefited from FEDER funds. ; Peer Reviewed

© 2014 Lozano et al. Recent studies in vitro have reported that the Methoprene-tolerant (Met) and Taiman (Tai) complex is the functional receptor of juvenile hormone (JH). Experiments in vivo of Met depletion have confirmed this factor's role in JH signal transduction, however, there is no equivalent data regarding Tai because its depletion in larval or nymphal stages of the beetle Tribolium castaneum and the bug Pyrrhocoris apterus results in 100% mortality. We have discovered that the cockroach Blattella germanica possesses four Tai isoforms resulting from the combination of two indels in the C-terminal region of the sequence. The presence of one equivalent indel-1 in Tai sequences in T. castaneum and other species suggests that Tai isoforms may be common in insects. Concomitant depletion of all four Tai isoforms in B. germanica resulted in 100% mortality, but when only the insertion 1 (IN-1) isoforms were depleted, mortality was significantly reduced and about half of the specimens experienced precocious adult development. This shows that Tai isoforms containing IN-1 are involved in transducing the JH signal that represses metamorphosis. Reporter assays indicated that both T. castaneum Tai isoforms, one that contains the IN-1 and another that does not (DEL-1) activated a JH response element (kJHRE) in Krüppel homolog 1 in conjunction with Met and JH. The results indicate that Tai is involved in the molecular mechanisms that repress metamorphosis, at least in B. germanica, and highlight the importance of distinguishing Tai isoforms when studying the functions of this transcription factor in development and other processes. ; Financial support for this research was provided by the Spanish MICINN (grant CGL2008-03517/BOS to XB and predoctoral fellowship to JL), Spanish MINECO (grant CGL2012-36251 to XB), from the Catalan Government (2009 SGR 1498), by the Japanese JSPS KAKENHI Grant Numbers, 2585023 to TK and 25252059 to TS, and the NIAS Strategic Research Fund. The research has also benefited from FEDER funds associated to Spanish grants. ; Peer Reviewed

The influence of DNA methylation on gene behavior and its consequent phenotypic effects appear to be very important, but the details are not well understood. Insects offer a diversity of DNA methylation modes, making them an excellent lineage for comparative analyses. However, functional studies have tended to focus on quite specialized holometabolan species, such as wasps, bees, beetles, and flies. Here, we have studied DNA methylation in the hemimetabolan insect Blattella germanica. In this cockroach, a gene involved in DNA methylation, DNA methyltransferase 1 (DNMT1), is expressed in early embryogenesis. In our experiments, RNAi of DNMT1 reduces DNA methylation and impairs blastoderm formation. Using reduced representation bisulfite sequencing and transcriptome analyses, we observed that methylated genes are associated with metabolism and are highly expressed, whereas unmethylated genes are related to signaling and show low expression. Moreover, methylated genes show greater expression change and less expression variability than unmethylated genes.Developmental Genetics; Molecular Biology ; This work was supported by the Spanish Ministry of Economy and Competitiveness (grants CGL2012- 36251, CGL2015-64727-P and PID2019-104483GB-I00 to XB, including FEDER funds), the CSIC (grant 2019AEP029), and the Catalan Government (grants 2014 SGR 619 and 2017 SGR 1030).

[Background] Argonaute proteins are key in RNA silencing. In Drosophila melanogaster, the five proteins of the Argonaute family participate in the pathways and mechanisms mediated by three types of small RNAs: piRNAs, miRNAs, and siRNAs. Two Argonaute proteins, Argonaute 1 (Ago1) and Argonaute 2 (Ago2), are associated with miRNA and siRNA mechanisms, which are the most thoroughly studied. The available data points to a sorting specialization of Ago1 for miRNAs and Ago2 for siRNAs. However, this has been demonstrated only in D. melanogaster, one of the most modified insects, which emerged some 100 million years ago. Thus, an important question is whether this association of Ago1 with miRNAs and Ago2 with siRNAs occurs generally in insects, or was a specific innovation in higher flies. ; [Methods] We addressed this question by using RNAi approaches and studying Ago1 and Ago2 functions in the German cockroach, Blattella germanica, a much less modified insect that emerged some 320 million years ago. ; [Results] The results showed that B. germanica does preferentially use Ago1 in the miRNA pathway, but can also use Ago2 in some cases. Conversely, Ago2 operates in the RNAi, in siRNA sorting, whereas Ago1 seems to have no relevant role in this process. ; [Conclusions and general significance] These basic associations are equivalent to those observed in D. melanogaster, implying that they have been evolutionary conserved from at least cockroach to flies, and possibly stem from the last common ancestor of extant insects. ; This work was supported by the Spanish Ministry of Economy and Competitiveness (grants CGL2012–36251 and CGL2015–64727-P to X.B., and CGL2016–76011-R to M.D.P. and J.L.M.) and the Catalan Government (2017 SGR 1030). It also received financial assistance from the European Fund for Economic and Regional Development (FEDER funds to X.B., M.D.P. and J.L.M.). ; Peer reviewed

The Piwi‐interacting RNA (piRNA) system is an evolutionarily conserved mechanism involved in the control of transposable elements and maintenance of genomic stability, especially in germ line cells and in early embryo stages. However, relevant particularities, both in mechanism and function, exist across species among metazoans and even within the insect class. As a member of the scarcely studied hemimetabolan group, Blattella germanica can be a suitable reference model to study insect evolution. We present the results of a stringent process of identification and study of expressed piRNAs for B. germanica across 11 developmental stages, ranging from unfertilized egg to nymphs and adult female. Our results confirm the dual origin of piRNA in this species, with a majority of them being generated from the primary pathway, and a smaller but highly expressed set of sequences participating in the secondary ("ping‐pong") reamplification pathway. An intriguing partial complementarity in expression is observed between the piRNA of the two biogenesis pathways, with those generated in the secondary pathway being quite restricted to early embryo stages. In addition, many piRNAs are exclusively expressed in late embryo and nymphal stages. These observations point at piRNA functions beyond the role of transposon control in early embryogenesis. Our work supports the view of a more complex scenario, with different sets of piRNAs acting in different times and having a range of functions wider than previously thought. ; This work was supported by the Spanish Ministry of Economy and Competitiveness (grants CGL2012-36251 and CGL2015-64727-P to X.B.; and CGL2016-76011-R to M.D.P.), the Spanish Ministry of Science and Innovation (grant BFU2011-22404) and the Catalan Government (2014 SGR 619). It also received financial assistance from the European Fund for Economic and Regional Development (FEDER funds to X.B. and M.D.P.). ; Peer reviewed

7 pages, 4 figures, supplementary Information accompanies this paper at http://dx.doi.org/10.1038/ncomms8814 ; Transmembrane glycerol transport is typically facilitated by aquaglyceroporins in Prokaryota and Eukaryota. In holometabolan insects however, aquaglyceroporins are absent, yet several species possess polyol permeable aquaporins. It thus remains unknown how glycerol transport evolved in the Holometabola. By combining phylogenetic and functional studies, here we show that a more efficient form of glycerol transporter related to the water-selective channel AQP4 specifically evolved and multiplied in the insect lineage, resulting in the replacement of the ancestral branch of aquaglyceroporins in holometabolan insects. To recapitulate this evolutionary process, we generate specific mutants in distantly related insect aquaporins and human AQP4 and show that a single mutation in the selectivity filter converted a water-selective channel into a glycerol transporter at the root of the crown clade of hexapod insects. Integration of phanerozoic climate models suggests that these events were associated with the emergence of complete metamorphosis and the unparalleled radiation of insects. © 2015 Macmillan Publishers Limited. All rights reserved ; This work was supported by the Research Council of Norway (RCN) project 204813/F20 to R.N.F., the Spanish Ministry of Economy and Competitiveness (MINECO), projects AGL2013-41196-R to J.C. and CGL2012-36251 to X.B. and the Catalan Government, grants 2014 SGR 1351 to J.C. and 2014 SGR 619 to X.B. The research has also benefitted from FEDER funds. F.C. and J.A.S. were supported by postdoctoral fellowships from the RCN (224816/E40 and 204813/F20, respectively) ; Peer Reviewed