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In this paper we explore high-throughput Illumina sequencing of nuclear protein-coding, ribosomal, and mitochondrial genes in small, dried insects stored in natural history collections. We sequenced one tenebrionid beetle and 12 carabid beetles ranging in size from 3.7 to 9.7 mm in length that have been stored in various museums for 4 to 84 years. Although we chose a number of old, small specimens for which we expected low sequence recovery, we successfully recovered at least some low-copy nuclear protein-coding genes from all specimens. For example, in one 56-year-old beetle, 4.4 mm in length, our de novo assembly recovered about 63% of approximately 41,900 nucleotides in a target suite of 67 nuclear protein-coding gene fragments, and 70% using a reference-based assembly. Even in the least successfully sequenced carabid specimen, reference-based assembly yielded fragments that were at least 50% of the target length for 34 of 67 nuclear protein-coding gene fragments. Exploration of alternative references for reference-based assembly revealed few signs of bias created by the reference. For all specimens we recovered almost complete copies of ribosomal and mitochondrial genes. We verified the general accuracy of the sequences through comparisons with sequences obtained from PCR and Sanger sequencing, including of conspecific, fresh specimens, and through phylogenetic analysis that tested the placement of sequences in predicted regions. A few possible inaccuracies in the sequences were detected, but these rarely affected the phylogenetic placement of the samples. Although our sample sizes are low, an exploratory regression study suggests that the dominant factor in predicting success at recovering nuclear protein-coding genes is a high number of Illumina reads, with success at PCR of COI and killing by immersion in ethanol being secondary factors; in analyses of only high-read samples, the primary significant explanatory variable was body length, with small beetles being more successfully sequenced.

A series of laboratory studies was conducted to assess the effect of droplet size on efficacy of pyrethrin aerosol against adults of Tribolium confusum Jacqueline DuVal, the confused flour beetle. A vertical flow aerosol exposure chamber that generated a standardized particle size diameter was used for these trials. In the first experiments, adults were exposed in the chamber for 2.5-45 min to aerosol dispensed at a volumetric median particle size diameter (VMD) of 16 mu m, and then held in the arenas in which they were exposed or transferred to new arenas with or without a flour food source. All adults were initially knocked down when removed from the chamber. Recovery from knockdown decreased as exposure interval increased, but the presence of a food source enhanced recovery at the lower exposure intervals. In the second experiment, the aerosol was applied at a VMD of 2 mu m and adults were exposed for between 5 and 75 min. Knockdown of adults was less than or equal to 10% when adults were removed from the chamber regardless of exposure time and afterward there was essentially complete recovery of adults. In the third and final experiment, the same 2- mu m VMD particle size and exposure times were used, but the concentration of aerosol was increased by approximately 4 compared with the previous experiment. In this test, initial knockdown was greater at the higher exposure intervals, but by 3 and 4 d posttreatment, recovery was again essentially 100%. This is the first published test assessing the efficacy of specific aerosol particle sizes on a stored product insect. Results indicate that particle size was a more important factor in conferring toxicity than the actual concentration or number of aerosol particles.

A review of genus-group names for darkling beetles in the family Tenebrionidae (Insecta: Coleoptera) is presented. A catalogue of 4122 nomenclaturally available genus-group names, representing 2307 valid genera (33 of which are extinct) and 761 valid subgenera, is given. For each name the author, date, page number, gender, type species, type fixation, current status, and first synonymy (when the name is a synonym) are provided. Genus-group names in this family are also recorded in a classification framework, along with data on the distribution of valid genera and subgenera within major biogeographical realms. A list of 535 unavailable genus-group names (e.g., incorrect subsequent spellings) is included. Notes on the date of publication of references cited herein are given, when known. The following genera and subgenera are made available for the first time: Anemiadena Bouchard & Bousquet, subgen. nov. (in Cheirodes Gené, 1839), Armigena Bouchard & Bousquet, subgen. nov. (in Nesogena Mäklin, 1863), Debeauxiella Bouchard & Bousquet, subgen. nov. (in Hyperops Eschscholtz, 1831), Hyperopsis Bouchard & Bousquet, subgen. nov. (in Hyperops Eschscholtz, 1831), Linio Bouchard & Bousquet, subgen. nov. (in Nilio Latreille, 1802), Matthewsotys Bouchard & Bousquet, gen. nov. , Neosolenopistoma Bouchard & Bousquet, subgen. nov. (in Eurynotus W. Kirby, 1819), Paragena Bouchard & Bousquet, subgen. nov. (in Nesogena Mäklin, 1863), Paulianaria Bouchard & Bousquet, gen. nov. , Phyllechus Bouchard & Bousquet, gen. nov. , Prorhytinota Bouchard & Bousquet, subgen. nov. (in Rhytinota Eschscholtz, 1831), Pseudorozonia Bouchard & Bousquet, subgen. nov. (in Rozonia Fairmaire, 1888), Pseudothinobatis Bouchard & Bousquet, gen. nov. , Rhytinopsis Bouchard & Bousquet, subgen. nov. (in Thalpophilodes Strand, 1942), Rhytistena Bouchard & Bousquet, subgen. nov. (in Rhytinota Eschscholtz, 1831), Spinosdara Bouchard & Bousquet, subgen. nov. (in Osdara Walker, 1858), Spongesmia Bouchard & Bousquet, subgen. nov. (in Adesmia Fischer, 1822), and Zambesmia Bouchard & Bousquet, subgen. nov. (in Adesmia Fischer, 1822). The names Adeps Gistel, 1857 and Adepsion Strand, 1917 syn. nov. [= Tetraphyllus Laporte & Brullé, 1831], Asyrmatus Canzoneri, 1959 syn. nov. [= Pystelops Gozis, 1910], Euzadenos Koch, 1956 syn. nov. [= Selenepistoma Dejean, 1834], Gondwanodilamus Kaszab, 1969 syn. nov. [= Conibius J.L. LeConte, 1851], Gyrinodes Fauvel, 1897 syn. nov. [= Nesotes Allard, 1876], Helopondrus Reitter, 1922 syn. nov. [= Horistelops Gozis, 1910], Hybonotus Dejean, 1834 syn. nov. [= Damatris Laporte, 1840], Iphthimera Reitter, 1916 syn. nov. [= Metriopus Solier, 1835], Lagriomima Pic, 1950 syn. nov. [= Neogria Borchmann, 1911], Orphelops Gozis, 1910 syn. nov. [= Nalassus Mulsant, 1854], Phymatium Billberg, 1820 syn. nov. [= Cryptochile Latreille, 1828], Prosoblapsia Skopin & Kaszab, 1978 syn. nov. [= Genoblaps Bauer, 1921], and Pseudopimelia Gebler, 1859 syn. nov. [= Lasiostola Dejean, 1834] are established as new synonyms (valid names in square brackets). Anachayus Bouchard & Bousquet, nom. nov. is proposed as a replacement name for Chatanayus Ardoin, 1957, Genateropa Bouchard & Bousquet, nom. nov. as a replacement name for Apterogena Ardoin, 1962, Hemipristula Bouchard & Bousquet, nom. nov. as a replacement name for Hemipristis Kolbe, 1903, Kochotella Bouchard & Bousquet, nom. nov. as a replacement name for Millotella Koch, 1962, Medvedevoblaps Bouchard & Bousquet, nom. nov. as a replacement name for Protoblaps G.S. Medvedev, 1998, and Subpterocoma Bouchard & Bousquet, nom. nov. is proposed as a replacement name for Pseudopimelia Motschulsky, 1860. Neoeutrapela Bousquet & Bouchard, 2013 is downgraded to a subgenus ( stat. nov. ) of Impressosora Pic, 1952. Anchomma J.L. LeConte, 1858 is placed in Stenosini: Dichillina (previously in Pimeliinae: Anepsiini); Entypodera Gerstaecker, 1871, Impressosora Pic, 1952 and Xanthalia Fairmaire, 1894 are placed in Lagriinae: Lagriini: Statirina (previously in Lagriinae: Lagriini: Lagriina); Loxostethus Triplehorn, 1962 is placed in Diaperinae: Diaperini: Diaperina (previously in Diaperinae: Diaperini: Adelinina); Periphanodes Gebien, 1943 is placed in Stenochiinae: Cnodalonini (previously in Tenebrioninae: Helopini); Zadenos Laporte, 1840 is downgraded to a subgenus ( stat. nov. ) of the older name Selenepistoma Dejean, 1834. The type species [placed in square brackets] of the following available genus-group names are designated for the first time: Allostrongylium Kolbe, 1896 [ Allostrongylium silvestre Kolbe, 1896], Auristira Borchmann, 1916 [ Auristira octocostata Borchmann, 1916], Blapidocampsia Pic, 1919 [ Campsia pallidipes Pic, 1918], Cerostena Solier, 1836 [ Cerostena deplanata Solier, 1836], Coracostira Fairmaire, 1899 [ Coracostira armipes Fairmaire, 1899], Dischidus Kolbe, 1886 [ Helops sinuatus Fabricius, 1801], Eccoptostoma Gebien, 1913 [ Taraxides ruficrus Fairmaire, 1894], Ellaemus Pascoe, 1866 [ Emcephalus submaculatus Brême, 1842], Epeurycaulus Kolbe, 1902 [ Epeurycaulus aldabricus Kolbe, 1902], Euschatia Solier, 1851 [ Euschatia proxima Solier, 1851], Heliocaes Bedel, 1906 [ Blaps emarginata Fabricius, 1792], Hemipristis Kolbe, 1903 [ Hemipristis ukamia Kolbe, 1903], Iphthimera Reitter, 1916 [ Stenocara ruficornis Solier, 1835], Isopedus Stein, 1877 [ Helops tenebrioides Germar, 1813], Malacova Fairmaire, 1898 [ Malacova bicolor Fairmaire, 1898], Modicodisema Pic, 1917 [ Disema subopaca Pic, 1912], Peltadesmia Kuntzen, 1916 [ Metriopus platynotus Gerstaecker, 1854], Phymatium Billberg, 1820 [ Pimelia maculata Fabricius, 1781], Podoces Péringuey, 1886 [ Podoces granosula Péringuey, 1886], Pseuduroplatopsis Pic, 1913 [ Borchmannia javana Pic, 1913], Pteraulus Solier, 1848 [ Pteraulus sulcatipennis Solier, 1848], Sciaca Solier, 1835 [ Hylithus disctinctus Solier, 1835], Sterces Champion, 1891 [ Sterces violaceipennis Champion, 1891] and Teremenes Carter, 1914 [ Tenebrio longipennis Hope, 1843]. Evidence suggests that some type species were misidentified. In these instances, information on the misidentification is provided and, in the following cases, the taxonomic species actually involved is fixed as the type species [placed in square brackets] following requirements in Article 70.3 of the International Code of Zoological Nomenclature: Accanthopus Dejean, 1821 [ Tenebrio velikensis Piller & Mitterpacher, 1783], Becvaramarygmus Masumoto, 1999 [ Dietysus nodicornis Gravely, 1915], Heterophaga Dejean, 1834 [ Opatrum laevigatum Fabricius, 1781], Laena Dejean, 1821, [ Scaurus viennensis Sturm, 1807], Margus Dejean, 1834 [ Colydium castaneum Herbst, 1797], Pachycera Eschscholtz, 1831 [ Tenebrio buprestoides Fabricius, 1781], Saragus Erichson, 1842 [ Celibe costata Solier, 1848], Stene Stephens, 1829 [ Colydium castaneum Herbst, 1797], Stenosis Herbst, 1799 [ Tagenia intermedia Solier, 1838] and Tentyriopsis Gebien, 1928 [ Tentyriopsis pertyi Gebien, 1940]. The following First Reviser actions are proposed to fix the precedence of names or nomenclatural acts (rejected name or act in square brackets): Stenosis ciliaris Gebien, 1920 as the type species for Afronosis G.S. Medvedev, 1995 [ Stenosis leontjevi G.S. Medvedev, 1995], Alienoplonyx Bremer, 2019 [ Alienolonyx ], Amblypteraca Mas-Peinado, Buckley, Ruiz & García-París, 2018 [ Amplypteraca ], Caenocrypticoides Kaszab, 1969 [ Caenocripticoides ], Deriles Motschulsky, 1872 [ Derilis ], Eccoptostira Borchmann, 1936 [ Ecoptostira ], † Eodromus Haupt, 1950 [† Edromus ], Eutelus Solier, 1843 [ Lutelus ], Euthriptera Reitter, 1893 [ Enthriptera ], Meglyphus Motschulsky, 1872 [ Megliphus ], Microtelopsis Koch, 1940 [ Extetranosis Koch, 1940, Hypermicrotelopsis Koch, 1940], Neandrosus Pic, 1921 [ Neoandrosus ], Nodosogylium Pic, 1951 [ Nodosogilium ], Notiolesthus Motschulsky, 1872 [ Notiolosthus ], Pseudeucyrtus Pic, 1916 [ Pseudocyrtus ], Pseudotrichoplatyscelis Kaszab, 1960 [ Pseudotrichoplatynoscelis and Pseudotrichoplatycelis ], Rhydimorpha Koch, 1943 [ Rhytimorpha ], Rhophobas Motschulsky, 1872 [ Rophobas ], Rhyssochiton Gray, 1831 [ Ryssocheton and Ryssochiton ], Sphaerotidius Kaszab, 1941 [ Spaerotidius ], Stira Agassiz, 1846 (Mollusca) [ Stira Agassiz, 1846 (Coleoptera)], Sulpiusoma Ferrer, 2006 [ Sulpiosoma ] and Taenobates Motschulsky, 1872 [ Taeniobates ]. Supporting evidence is provided for the conservation of usage of Cyphaleus Westwood, 1841 nomen protectum over Chrysobalus Boisduval, 1835 nomen oblitum.

Phylogenetic trees in insects are frequently dated by applying a “standard” mitochondrial DNA (mtDNA) clock estimated at 2.3% My−1, but despite its wide use reliable calibration points have been lacking. Here, we used a well-established biogeographic barrier, the mid-Aegean trench separating the western and eastern Aegean archipelago, to estimate substitution rates in tenebrionid beetles. Cytochrome oxidase I (cox1) for six codistributed genera across 28 islands (444 individuals) on both sides of the mid-Aegean trench revealed 60 independently coalescing entities delimited with a mixed Yule-coalescent model. One representative per entity was used for phylogenetic analysis of mitochondrial (cox1, 16S rRNA) and nuclear (Mp20, 28S rRNA) genes. Six nodes marked geographically congruent east–west splits whose separation was largely contemporaneous and likely to reflect the formation of the mid-Aegean trench at 9–12 Mya. Based on these “known” dates, a divergence rate of 3.54% My−1 for the cox1 gene (2.69% when combined with the 16S rRNA gene) was obtained under the preferred partitioning scheme and substitution model selected using Bayes factors. An extensive survey suggests that discrepancies in mtDNA substitution rates in the entomological literature can be attributed to the use of different substitution models, the use of different mitochondrial gene regions, mixing of intraspecific with interspecific data, and not accounting for variance in coalescent times or postseparation gene flow. Different treatments of these factors in the literature confound estimates of mtDNA substitution rates in opposing directions and obscure lineage-specific differences in rates when comparing data from various sources.

The fossil record of Tenebrionidae (excluding the Quartenary) is presented. In total, 122 fossil species, clearly belonging to the family, are known; some beetles were determined only to genus; 78 genera are listed in the fossil record, including 29 extinct genera. The great diversity of tenebrionids occurs in the Lower Cretaceous Lagerstätte of China (Yixian Formation), Middle Paleocene of France (Menat), Lower Eocene deposits of Germany (Geiseltal), Upper Eocene Baltic amber (Eastern Europe), Upper Eocene deposits of Florissant Formation (USA) and Miocene (Dominican amber). Tenebrionids of the following major lineages, including seven subfamilies, are currently known in the fossil record. These include the lagrioid branch (Lagriinae, Nilioninae), pimelioid branch (Pimeliinae), and tenebrioid branch (Alleculinae, Tenebrioninae, Diaperinae, Stenochiinae). The importance of the fossil record for evolutionary reconstructions and phylogenetic patterns is discussed. The oldest Jurassic and Early Cretaceous darkling beetles of the tenebrionoid branch consist of humid-adapted groups from the extant tribes Alleculini, Ctenopodiini (Alleculinae), and Alphitobiini (Tenebrioninae). Thus, paleontological evidence suggests that differentiation of the family started at least by the Middle Jurassic but does not indicate that xerophilic darkling beetles differentiated much earlier than mesophilic groups.

The effect of spinosad exposure on the susceptibility of pyrethroid- and organophosphate-resistant populations of lesser mealworm, Alphitobius diaperinus (Panzer) (Coleoptera: Tenebrionidae), to insecticides was investigated in broiler house farm and laboratory studies. A field pyrethroid- and organophosphate-resistant population showed a 3.6-fold increase in susceptibility to gamma -cyhalothrin following spinosad treatment. Overall, cyfluthrin- and fenitrothion-resistant field populations were more susceptible to these insecticides following spinosad treatments, but populations that were not resistant showed no change in susceptibility following spinosad treatment. In a related study, three broiler farm beetle populations with very similar levels of cyfluthrin and gamma -cyhalothrin resistance and similar susceptibilities to spinosad were used to investigate temporal effects of spinosad field treatments on the susceptibility to pyrethroids. Farm insecticide regimes applied at the start of each flock differed: the control broiler house received no insecticide applications, another house was systematically treated with cyfluthrin at the start of each study flock, and the third house was systematically treated with spinosad at the start of five flocks. Afterwards, treatments reverted to cyfluthrin on all farms. At the end of flocks, beetles were tested with cyfluthrin, gamma -cyhalothrin, and spinosad. The control and cyfluthrin house beetles did not change susceptibility to pyrethroids over the period of the study. In the spinosad house, spinosad had no effect on spinosad susceptibility but dramatically increased cyfluthrin and gamma -cyhalothrin susceptibilities. These new susceptibilities were maintained while spinosad applications continued, but pyrethroid susceptibility declined once spinosad applications ceased. This study provides evidence of a synergistic interaction between spinosad and pyrethroids in pyrethroid-resistant beetles. This evidence has significant implications for management of insecticide-resistant populations through an integrated spinosad-pyrethroid strategy that aims to minimize insecticide use while enhancing control.

Most insecticides commonly used in storage facilities are synthetic, an issue that generates concerns about food safety and public health. Therefore, the development of eco-friendly pest management tools is urgently needed. In the present study, a 6% (w/w) Hazomalania voyronii essential oil-based nanoemulsion (HvNE) was developed and evaluated for managing Tribolium confusum, T. castaneum, and Tenebrio molitor, as an eco-friendly wheat protectant. Larval and adult mortality was evaluated after 4, 8, and 16 h, and 1, 2, 3, 4, 5, 6, and 7 days, testing two HvNE concentrations (500 ppm and 1000 ppm). T. confusum and T. castaneum adults and T. molitor larvae were tolerant to both concentrations of the HvNE, reaching 13.0%, 18.7%, and 10.3% mortality, respectively, at 1000 ppm after 7 days of exposure. However, testing HvNE at 1000 ppm, the mortality of T. confusum and T. castaneum larvae and T. molitor adults 7 days post-exposure reached 92.1%, 97.4%, and 100.0%, respectively. Overall, the HvNE can be considered as an effective adulticide or larvicide, depending on the target species. Our results highlight the potential of H. voyronii essential oil for developing green nanoinsecticides to be used in real-world conditions against key stored-product pests.

The aim of this review is to compile up-to-date information on the superworm, Zophobas morio (F.), regarding its biology and ecology, but also its further potential for use as a nutrient source for food and feed. We illustrate certain basic characteristics of the morphology and bio-ecology of this species, which is marginally considered as a ‘pest’ in durable amylaceous commodities. More recent data show that Z. morio can be a valuable nutrient and antimicrobial source that could be utilized further in insect-based feed and food production. The inclusion of this species in aquafeed has provided promising results in a wide range of feeding trials, both in terms of fish development and health. Additional data illustrate its potential for use in poultry, indicating that this species provides comparable results with those of other insect species that are used in feed. Moreover, Z. morio can be a viable waste management agent. This review aims to summarize the available data and underline data gaps for future research, toward the potential of the utilization of Z. morio for human food and animal feed. Based on the data presented, Z. morio appears to be a well-promising insect-based protein source, which potential still remains to be unfold.

The evolutionary success of beetles and numerous other terrestrial insects is generally attributed to co-radiation with flowering plants but most studies have focused on herbivorous or pollinating insects. Non-herbivores represent a significant proportion of beetle diversity yet potential factors that influence their diversification have been largely unexamined. In the present study, we examine the factors driving diversification within the Scarabaeidae, a speciose beetle family with a range of both herbivorous and non-herbivorous ecologies. In particular, it has been long debated whether the key event in the evolution of dung beetles (Scarabaeidae: Scarabaeinae) was an adaptation to feeding on dinosaur or mammalian dung. Here we present molecular evidence to show that the origin of dung beetles occurred in the middle of the Cretaceous, likely in association with dinosaur dung, but more surprisingly the timing is consistent with the rise of the angiosperms. We hypothesize that the switch in dinosaur diet to incorporate more nutritious and less fibrous angiosperm foliage provided a palatable dung source that ultimately created a new niche for diversification. Given the well-accepted mass extinction of non-avian dinosaurs at the Cretaceous-Paleogene boundary, we examine a potential co-extinction of dung beetles due to the loss of an important evolutionary resource, i.e., dinosaur dung. The biogeography of dung beetles is also examined to explore the previously proposed \"out of Africa\" hypothesis. Given the inferred age of Scarabaeinae as originating in the Lower Cretaceous, the major radiation of dung feeders prior to the Cenomanian, and the early divergence of both African and Gondwanan lineages, we hypothesise that that faunal exchange between Africa and Gondwanaland occurred during the earliest evolution of the Scarabaeinae. Therefore we propose that both Gondwanan vicariance and dispersal of African lineages is responsible for present day distribution of scarabaeine dung beetles and provide examples.

Traps baited with pheromones are used to monitor the red flour beetle, Tribolium castaneum (Herbst) (Coleoptera: Tenebrionidae), populations in flour mills to aid in making pest management decisions, but the factors that influence T. castaneum movement are not fully understood. We investigated the impact of photoperiod, light intensity, temperature, and relative humidity on flight initiation. The percentage of adults initiating flight reached a maximum at 30-35 degree C, and then fell to zero at 22.5 and 45 degree C. Only 2% of beetles flew in complete darkness, and the number of beetles initiating flight increased to 41% under 18 h of light and then decreased slightly to 37% under 24 h of light. Rates of flight initiation did not vary with light intensities from 1,784 to 4,356 lux or relative humidities from 25 to 85%. Thus, temperature and photoperiod are the main abiotic factors tested that impact flight initiation in T. castaneum, which have broad ranges of temperatures and photoperiods over which they can fly. The current results should be useful in helping to interpret trap catches based on abiotic conditions during the trapping period, and the results should be useful in helping to understand T. castaneum movement outside grain storages and processing facilities and their potential to infest structures.