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Abstract

Insects are one of the few groups of animals that developed the ability of active flight. Such mobility allowed the group to successfully explore and thrive in nearly all kinds of ecological niches. At the same time, during the evolutionary history of insects, due to high costs of wing development, flight was lost independently in many groups. In beetles, the reduction or complete loss of hind wings has been reported in multiple lineages, especially in several extreme paedomorphic and larviform females, mainly in Elateroidea, in which not only the hind wings but also the elytra are lost. However, the complete absence of elytra in adult males was hitherto unknown, despite nearly half a million described species in Coleoptera. In this study, we report the discovery of Xenomorphon baranowskii gen. et sp. nov., the first completely anelytrous and wingless adult male beetle, belonging to the family Lycidae (Coleoptera: Elateroidea). Xenomorphon baranowskii is illustrated, described, and provisionally placed in Calopterini, based on our morphology-based phylogenetic analyses. We discuss the possible scenarios that could lead to such a rare event, when a beetle loses its elytra, and its evolutionary consequences.

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... The morphological matrix is based on that of Bocakova (2005), updated by Nascimento et al. (2020) and Ferreira et al. (2023). The dataset was expanded by the inclusion of two recently proposed Eurrhacini genera (Atlanticolycus, Cladocalon) and the new one described here (altogether six newly coded species). ...
... Support for a monophyletic origin of Calopterini and the subtribe Calopterina has been confirmed by previous (Bocakova 2005;Nascimento et al. 2020;Ferreira et al. 2023) and our current (this study) morphology-based analyses. However, the formerly recovered Acroleptina, comprising all neotenic calopterins, is now predominantly paraphyletic (Table 2) and split into two lineages. ...
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The first phylogenetic inference of Calopterini and Eurrhacini focused on Calocladon and related taxa was carried out. A data matrix composed of 46 species and 51 morphological characters was assembled and analyzed using parsimony and model-based approaches. Eurrhacini were recovered monophyletic. Furthermore, phylogenetic analyses highly supported the Calocladon clade including also Atlanticolycus, Cladocalon, and Gorhamiumgen. nov. as its sister clade. Our trees consistently recovered monophyly of the new genus with two new species: Gorhamium bidentatumsp. nov. (Panama, Baru Volcano) and G. unidentatumsp. nov. from the Pacific slopes of Ecuador. A revised key to the genera of Eurrhacini is given and illustrations of distinguishing characters are provided. Phylogenetic relationships of Eurrhacini and character evolution are discussed.
... In some older cladistic studies, autapomorphies were often neglected and altogether excluded, as they did not influence at all the topology of a tree generated via maximum parsimony (Kitching et al. 1998). We briefly present some arguments in favor of always including autapomorphic characters; see Yeates (1992) for a more complete argument: 1) autapomorphies have the potential to be diagnostic characters to terminals (higher taxa, and species), therefore, they have taxonomic importance (e.g., Ferreira et al. 2023); 2) it is well-known that the consistency index (c or ci) is "inflated" by the inclusion of autapomorphies (Brooks et al. 1986, Carpenter 1988), but there is no need to exclude them since most popular phylogenetic software, e.g., TNT 1.5 (Goloboff and Catalano 2016) and PAUP* 4.0 (Swofford and Bell 2017), report the consistency index both with and without autapo-morphies (so the inclusion of autapomorphies is not detrimental in any point and also does not significantly increase tree search time); 3) in maximum likelihood (Felsenstein 1973(Felsenstein , 1981 and Bayesian Yang 1996, Yang andRannala 1997) approaches, autapomorphies inform branch lengths and equilibrium state frequencies, moreover, in the context of these methods, autapomorphies are phylogenetically informative (Felsenstein 1978, Lewis 2001. ...
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... This is a popular approach (e.g. Buckley 2002, Aris-Brosou 2003, Planet 2006, Ferreira et al. 2023) also applied in phylogenomic studies (e.g. Cui et al. 2013, Arcila et al. 2017, Díaz-Escandón et al. 2022, Dietz et al. 2023, including anchored hybrid enrichment datasets (e.g. ...
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... Several features found in Penicillophorinae were putatively associated with the miniaturization process in other Phengodidae taxa (e.g., Roza et al. 2017): reduction in the number of antennomeres, reduction or fusion of antennal flabellae (what in Penicillophorinae might be interpreted as extreme reduction or loss of antennal flabellae), and reduction of wing venation. These characteristics are known to occur in several small-bodied phengodid genera, such as Akamboja Roza et al., 2017, Cleidella Roza and Mermudes, 2020, Decamastinocerus Wittmer, 1988, Euryognathus Wittmer, 1976, Microcydistus Kundrata et al., 2021, Microphrixothrix Roza and Mermudes, 2019, Steneuryopa Wittmer, 1986 In this context, most unique characteristics of Penicillophorinae, such as simple antennae without flabellae, reduction in palpomere number, and reduction of wing venation, could actually be homoplastic products of independent miniaturization processes and the effects of the paedomorphic syndrome (e.g., Bocak et al. 2016;Ferreira et al. , 2023Kundrata et al. 2015Kundrata and Bocak 2019;Tokareva et al. 2023). In addition, some characters might be a misinterpretation of broken or poorly preserved specimens (such as the antennomere number in Adendrocera). ...
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The Penicillophorinae (Phengodidae) are a rather morphologically modified and enigmatic group of railroad-worm beetles. Specimens are rarely collected, and nearly all species are only known from their holotypes. A recent trip by AKH to Guatemala resulted in the collection of a specimen of Adendrocera Wittmer, 1976, the second ever known to be collected for this genus. Further studies of that individual indicated that it was a new species. In this paper, we describe Adendrocera carmelita Roza, Hansen, and Ferreira, new species and redescribe Adendrocera flavula Wittmer, 1976, the type species of the genus. We note that Adendrocera is of feminine gender and correct the inflection of the adjective constituting the type species epithet flavulum (neuter) to flavula (feminine). We also discuss the hypothesis that the convergent morphology within this subfamily might be a result of miniaturization and paedomorphic processes.
... The Eurrhacini is a monophyletic group (Bocakova 2005, Nascimento et al. 2020, Masek et al 2018, Ferreira et al. 2023) currently comprised of 8 genera and 123 species, known to be distributed in almost all Neotropical countries (Kleine 1933, Blackwelder 1945, Bocakova 2003, Nascimento & Bocakova 2017, 2019, 2022, Nascimento et al. 2020, Nascimento 2023. For more than a century Calocladon was only known from Central America, but more recently it has been recognized to have a wider distribution in the Neotropics (Constantin 2010, Nascimento 2013. ...
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Atlanticolycus gen. nov., is herein described as a new genus of Neotropical Lycidae in the Eurrhacini tribe. Five new species are described: Atlanticolycus camposgerais sp. nov., A. japi sp. nov., A. itatiaia sp. nov., A. ilhabela sp. nov., and A. morretes sp. nov., all distributed in Brazilian Atlantic Forest. Illustrations of diagnostic characters, a key for species identification and a key to genera of Eurrhacini with flabellate antennae are presented.
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External and internal structures of the thorax of the myrmecophile beetle Claviger testaceus (Clavigeritae, Pselaphinae) were examined and documented with state-of-the-art visualization techniques. Following a general trend in the omaliine lineage (Staphylinidae), the skeletal elements of the pro- and pterothorax in Claviger reach a maximum degree of compactness, with largely reduced inter- and intrasegmental sutures and skeletal elements linked with the flight apparatus. The musculature, especially metathoracic direct and indirect flight muscles, also shows a high degree of reduction. Two forms of wings were found among individuals of C. testaceus , both non-functional and representing an advanced stage of reduction. However, that wing vestiges are still present and the metanotum, only slightly reduced, suggests that loss of flight in this species is likely the result of a young evolutionary process. Several structures are linked with myrmecophilous habits: small body size facilitates transportation of beetles by ant workers and makes it easier to move inside nest tunnels; the remarkably compact body and mechanically robust appendages make the beetles less vulnerable to attacks by ant mandibles; the improved elytral interlocking mechanism and unusually expanded epipleura enhance the protection of vulnerable dorsal parts of the pterothorax and anterior abdomen; and glands associated with trichomes on the posterolateral elytral angle produce secretions attractive for ants. Various modifications of the thorax and anterior abdomen lead to an optimization of intimate associations with ants. The morphological syndrome enabling these beetles to cope with life in ant colonies evolved in several steps. This is suggested by an increasing solidification of the thoracic skeleton in related non-myrmecophilous groups and also by less modified related clavigerites;for instance, ant-associated tropical species are still able to fly.
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Cafius gigas Lea, 1929 (Coleoptera: Staphylinidae) was a large rove beetle endemic to Lord Howe Island (LHI) resembling Cafius and the LHI flightless endemic Hesperus dolichoderes (Lea, 1925). Like several other LHI endemics, C. gigas became extinct due to human-introduced rats. It is a legacy species valuable for understanding the LHI biota in terms of evolutionary biology and historical biogeography. Whether C. gigas was a member of Cafius Curtis, 1829, restricted to oceanic shores and prone to trans-oceanic dispersal, or related to H. dolichoderes, would have different implications. We subjected C. gigas to a total-evidence phylogenetic analyses of morphological and molecular data using model-based and parsimony methods. As a result, it is transferred to Hesperus Fauvel, 1874 with the new combination Hesperus gigas (Lea, 1929) comb. nov. Our analysis indicates that the montane leaf litter inhabitant H. gigas evolved neither in situ nor from a seashore Cafius-ancestor, or from an ancestor shared by two other LHI endemic congeners, Hesperus pacificus Olliff, 1887 and H. dolichoderes. It also suggests that all three Hesperus species that currently occur on LHI could have evolved on various seamounts at various times before reaching LHI.
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• Bats (order Chiroptera) are the only mammals capable of powered flight, and this may be an important factor behind their rapid diversification into the over 1400 species that exist today – around a quarter of all mammalian species. Though flight in bats has been extensively studied, the evolutionary history of the ability to fly in the chiropterans remains unclear. • We provide an updated synthesis of current understanding of the mechanics of flight in bats (from skeleton to metabolism), its relation to echolocation, and where previously articulated evolutionary hypotheses for the development of flight in bats stand following recent empirical advances. We consider the gliding model, and the echolocation‐first, flight‐first, tandem development, and diurnal frugivore hypotheses. In the light of the recently published description of the web‐winged dinosaur Ambopteryx longibrachium, we draw together all the current evidence into a novel hypothesis. • We present the interdigital webbing hypothesis: the ancestral bat exhibited interdigital webbing prior to powered flight ability, and the Yangochiroptera, Pteropodidae, and Rhinolophoidea evolved into their current forms along parallel trajectories from this common ancestor. Thus, we suggest that powered flight may have evolved multiple times within the Chiroptera and that similarity in wing morphology in different lineages is driven by convergence from a common ancestor with interdigital webbing.
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Cafius gigas Lea, 1929 (Coleoptera: Staphylinidae) was a large rove beetle endemic to Lord Howe Island (LHI) resembling Cafius and the LHI flightless endemic Hesperus dolichoderes (Lea, 1925). Like several other LHI endemics, C. gigas became extinct due to human-introduced rats. It is a legacy species valuable for understanding the LHI biota in terms of evolutionary biology and historical biogeography. Whether C. gigas was a member of Cafius Curtis, 1829, restricted to oceanic shores and prone to trans-oceanic dispersal, or related to H. dolichoderes, would have different implications. We subjected C. gigas to a total-evidence phylogenetic analyses of morphological and molecular data using model-based and parsimony methods. As a result, it is transferred to Hesperus Fauvel, 1874 with the new combination Hesperus gigas (Lea, 1929) comb. nov. Our analysis indicates that the montane leaf litter inhabitant H. gigas evolved neither in situ nor from a seashore Cafius-ancestor, or from an ancestor shared by two other LHI endemic congeners, Hesperus pacificus Olliff, 1887 and H. dolichoderes. It also suggests that all three Hesperus species that currently occur on LHI could have evolved on various seamounts at various times before reaching LHI.
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The Neotropical lycid genus Acroleptus Bourgeois, 1886 is revised and illustrations of diagnostic characters, geographic distribution maps and an identification key to the species are presented. Acroleptus chevrolati Bourgeois, 1886 is redescribed and a lectotype and a paralectotype are designated; two new species are described for the genus: Acroleptus alvarengai new species and Acroleptus limai new species. Two Aporrhipis Pascoe, 1887 species are described: Aporrhipis obrieni new species and Aporrhipis milleri new species and a key and geographic distribution map to the species of the genus are provided. Acroleptus costae Ferreira, 2015 is moved to the genus Paracroleptus new genus, a new genus erected to accommodate Paracroleptus costae (Ferreira, 2015) new combination. http://www.zoobank.org/urn:lsid:zoobank.org:pub:5F72E2BB-48BF-491D-8D3A-E7D878222B72
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Lycidae are among the better studied groups in the superfamily Elateroidea, however despite the progress in the taxonomic understanding of the Neotropical fauna, much still remains unknown and undescribed in the region. The description of the new genus Xenolycus gen. nov., from Serra dos Órgãos, a subrange of the Serra do Mar mountain range, in the Atlantic Rainforest in Rio de Janeiro State, Brazil, contributes to the knowledge of the Neotropical Lycidae fauna. The new genus can be distinguished from all other known Calopterini and Neotropical Lycidae by the combination of a pronotum with a wide, deep and strongly visible longitudinal cell in the disc area, the filiform antennae, the dehiscent elytra with reticulation strongly reduced and bearing only two weakly developed elytral costae and the mouthparts partially reduced, with rudimentary, barely visible mandibles. The type species, Xenolycus costae sp. nov., is illustrated and diagnostic characters and a discussion on the tribal placement of the new genus are provided.
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Fireflies (Lampyridae Rafinesque) are a diverse family of beetles which exhibit an array of morphologies including varying antennal and photic organ features. Due in part to their morphological diversity, the classification within the Lampyridae has long been in flux. Here we use an anchored hybrid enrichment approach to reconstruct the most extensive molecular phylogeny of Lampyridae to date (436 loci and 98 taxa) and use this phylogeny to evaluate the higher-level classification of the group. None of the currently recognized subfamilies were recovered as monophyletic with high support. We propose several classification changes supported by both phylogenetic and morphological evidence: 1) Pollaclasis Newman, Vestini McDermott (incl. Vesta Laporte, Dodacles Olivier, Dryptelytra Laporte, and Ledocas Olivier), Photoctus McDermott, and Araucariocladus Silveira & Mermudes are transferred to Lampyridae incertae sedis, 2) Psilocladinae Mcdermott, 1964status novum is reestablished for the genus Psilocladus Blanchard, 3) Lamprohizini Kazantsev, 2010 is elevated to Lamprohizinae Kazantsev, 2010status novum and Phausis LeConte is transferred to Lamprohizinae, 4) Memoan Silveira and Mermudes is transferred to Amydetinae Olivier, and 5) Scissicauda McDermott is transferred to Lampyrinae Rafinesque.
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The order Coleoptera (beetles) is arguably the most speciose group of animals, but the evolutionary history of beetles, including the impacts of plant feeding (herbivory) on beetle diversification, remain poorly understood. We inferred the phylogeny of beetles using 4,818 genes for 146 species, estimated timing and rates of beetle diversification using 89 genes for 521 species representing all major lineages and traced the evolution of beetle genes enabling symbiont-independent digestion of lignocellulose using 154 genomes or transcriptomes. Phylogenomic analyses of these uniquely comprehensive datasets resolved previously controversial beetle relationships, dated the origin of Coleoptera to the Carboniferous, and supported the codiversification of beetles and angiosperms. Moreover, plant cell wall-degrading enzymes (PCWDEs) obtained from bacteria and fungi via horizontal gene transfers may have been key to the Mesozoic diversification of herbivorous beetles—remarkably, both major independent origins of specialized herbivory in beetles coincide with the first appearances of an arsenal of PCWDEs encoded in their genomes. Furthermore, corresponding (Jurassic) diversification rate increases suggest that these novel genes triggered adaptive radiations that resulted in nearly half of all living beetle species. We propose that PCWDEs enabled efficient digestion of plant tissues, including lignocellulose in cell walls, facilitating the evolution of uniquely specialized plant-feeding habits, such as leaf mining and stem and wood boring. Beetle diversity thus appears to have resulted from multiple factors, including low extinction rates over a long evolutionary history, codiversification with angiosperms, and adaptive radiations of specialized herbivorous beetles following convergent horizontal transfers of microbial genes encoding PCWDEs.
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Phasmatodea comprises over 3,000 extant species and stands out as one of the last remaining insect orders for which a robust, higher-level phylogenetic hypothesis is lacking. New research suggests that the extant diversity is the result of a surprisingly recent and rapid radiation that has been difficult to resolve with standard Sanger sequence data. In order to resolve the early branching events of stick and leaf insects, we analyzed transcriptomes from 61 species, including 38 Phasmatodea species comprising all major clades and 23 outgroup taxa, including all other Polyneoptera orders. Using a custom-made ortholog set based on reference genomes from four species, we identified on average 2,274 orthologous genes in the sequenced transcriptomes. We generated various sub-alignments and performed maximum-likelihood analyses on several representative datasets to evaluate the effect of missing data and matrix composition on our phylogenetic estimates. Based on our new data, we are able to reliably resolve the deeper nodes between the principal lineages of extant Phasmatodea. Among Euphasmatodea, we provide strong evidence for a basal dichotomy of Aschiphasmatodea and all remaining euphasmatodeans, the Neophasmatodea. Within the latter clade, we recovered a previously unrecognized major New World and Old World lineage, for which we introduce the new names Oriophasmata tax. nov. (“Eastern phasmids”) and Occidophasmata tax. nov. (“Western phasmids”). Occidophasmata comprise Diapheromerinae, Pseudophasmatinae, and Agathemera, whereas all remaining lineages form the Oriophasmata, including Heteropterygidae, Phylliinae, Bacillus, Lonchodidae (Necrosciinae + Lonchodinae), Clitumninae, Cladomorphinae, and Lanceocercata. We furthermore performed a divergence time analysis and reconstructed the historical biogeography for stick and leaf insects. Phasmatodea either originated in Southeast Asia or in the New World. Our results suggest that the extant distribution of Phasmatodea is largely the result of dispersal events in a recently and rapidly diversified insect lineage rather than the result of vicariant processes.
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Since the description of the genus Cheguevaria as incertae sedis (Lampyridae), the placement of these beetles has been uncertain. This study is the first to address the placement of this genus based on a molecular phylogenetic analysis. We used three genes (18S rRNA, rrnl mitochondrial DNA and cox1 mitochondrial DNA) and a maximum likelihood approach with W-IQ-TREE to support Cheguevaria as a member of the Lampyridae and recognize it as the sole genus in the new subfamily Cheguevariinae stat. nov.
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Se describe Lycomesus llorentei gen. nov., sp. nov. (Coleoptera: Lycidae: Lycini), procedente de San Luis Potosí, México. Se presenta un cuadro comparativo con los caracteres diagnósticos entre los miembros de Lycini en Norteamérica: Lycus, Lycostomus, Lyconotus y Lycomesus.
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Flight loss in birds is as characteristic of the class Aves as flight itself. Although morphological and physiological differences are recognized in flight-degenerate bird species, their contributions to recurrent flight degeneration events across modern birds and underlying genetic mechanisms remain unclear. Here, in an analysis of 295 million nucleotides from 48 bird genomes, we identify two convergent sites causing amino acid changes in ATGLSer321Gly and ACOT7Ala197Val in flight-degenerate birds, which to our knowledge have not previously been implicated in loss of flight. Functional assays suggest that Ser321Gly reduces lipid hydrolytic ability of ATGL, and Ala197Val enhances acyl-CoA hydrolytic activity of ACOT7. Modeling simulations suggest a switch of main energy sources from lipids to carbohydrates in flight-degenerate birds. Our results thus suggest that physiological convergence plays an important role in flight degeneration, and anatomical convergence often invoked may not.
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The mesosomal skeletomuscular system of workers of Myrmecia nigrocincta was examined. A broad spectrum of methods was used, including micro-computed tomography combined with computer-based 3D reconstruction. An optimized combination of advanced techniques not only accelerates the acquisition of high quality anatomical data, but also facilitates a very detailed documentation and vi- sualization. This includes fine surface details, complex configurations of sclerites, and also internal soft parts, for instance muscles with their precise insertion sites. Myrmeciinae have arguably retained a number of plesiomorphic mesosomal features, even though recent mo- lecular phylogenies do not place them close to the root of ants. Our mapping analyses based on previous morphological studies and recent phylogenies revealed few mesosomal apomorphies linking formicid subgroups. Only five apomorphies were retrieved for the family, and interestingly three of them are missing in Myrmeciinae. Nevertheless, it is apparent that profound mesosomal transformations took place in the early evolution of ants, especially in the flightless workers. The modified mesosoma is characterized by four character complexes: a) an enlarged prothorax with elongate procoxae, a large plate-like pronotum, strongly developed muscles of the forelegs and especially of the neck region; b) highly differentiated legs with complex cleaning and attachment devices; c) a reduced flight apparatus with greatly simplified pterothoracic musculature and mechanically reinforced exoskeleton and d) strongly developed specialized muscles inserted on the base of the metasoma. Structural modifications of the prothorax and neck region allow ant workers to transport items efficiently with a highly movable head with strongly developed cervical muscles. Their differentiated legs enable them to move efficiently on various surfaces and to maintain their complex apparatus of sensilla. The mechanically reinforced mesosoma provides protection against predators and likely against detrimental environmental agents. The enhanced movability of the metasoma increases the defensive capacity with a sting or other mechanisms.
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A review of Lucaina Dugès, 1879 with illustrations of diagnostic characters, distribution maps, and an identification key to the species is presented. Lucaina schini Dugès, 1879, Lucaina discoidalis Horn, 1885, and Lucaina marginata Gorham, 1884 are redescribed, and their status as valid species is confirmed. Macrolygistopterus bajacalifornicus (Zaragoza, 2003) is transferred to Lucaina bajacalifornica (Zaragoza, 2003), new combination. Lucaina greeni Ferreira and Ivie, new species and Lucaina milleri Ferreira and Ivie, new species are described. Lectotypes and paralectotypes are designated for L. marginata Gorham, 1884, L. schini Dugès, 1879, and L. discoidalis Horn, 1885.
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We coin the term "taxon expeditions" for citizen scientists' field courses to carry out publishable taxonomic work in close association with trained taxonomists. During the first-ever taxon expedition, in Maliau Basin Studies Centre, Sabah, Malaysian Borneo, the participants sampled leaf litter beetles from lowland dipterocarp forest using the Winkler apparatus. The collected material proved to contain at least three undescribed species of small-bodied (ca. 1 mm long) hemispherical litter-dwelling Coleoptera. As part of the field course work, taxonomic descriptions were prepared for the chrysomelid Clavicornaltica sabahensis sp. n. and the leiodids Colenisia chungi sp. n. and Dermatohomoeus maliauensis sp. n.
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The standard bootstrap (SBS), despite being computationally intensive, is widely used in maximum likelihood phylogenetic analyses. We recently proposed the ultrafast bootstrap approximation (UFBoot) to reduce computing time while achieving more unbiased branch supports than SBS under mild model violations. UFBoot has been steadily adopted as an efficient alternative to SBS and other bootstrap approaches. Here, we present UFBoot2, which substantially accelerates UFBoot and reduces the risk of overestimating branch supports due to polytomies or severe model violations. Additionally, UFBoot2 provides suitable bootstrap resampling strategies for phylogenomic data. UFBoot2 is 778 times (median) faster than SBS and 8.4 times (median) faster than RAxML rapid bootstrap on tested datasets. UFBoot2 is implemented in the IQ-TREE software package version 1.6 and freely available at http://www.iqtree.org.
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Book
We share the earth with a wide variety of animal species, each of which brings something special to the diversity of the planet. By knowing more about how animals behave and live, we gain a greater understanding of how life evolved and the importance of biodiversity. This volume provides a complete guide to those birds that have evolved a trait that would seem to harm their ability to survive - flightlessness. Flight has its advantages - why would some birds be flightless?Flightless Birdscovers the loss of flight in birds, both permanently after years of evolution, and temporarily as a result of unusual molting behavior, and those species that are in various stages of losing their flight. The book provides a thorough guide, perfect for research papers in biology classes, for understanding the behavior and biodiversity of a fascinating and unusual group of animals. Flightless Birdsincludes sections on the major groups of flightless birds: Rarities whose ancient ancestors were on the continents when they broke away millions of years ago, and who survived despite competing with mammals; birds that were marooned on islands in the ocean, where food was plentiful and predators absent; penguins, which evolved alongside seas teeming with food and had no need to fly, and the special case of New Zealand's many flightless species which evolved in a predator-free paradise but could not cope with the settlers and their alien animals; and the many species which have become extinct within historic times. Beautifully illustrated, with numerous color photos,Flightless Birdsprovides copious material for understanding these unusual animals.
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The Leptolycini are a group of Lycidae endemic to the West Indies. Leptolycini adult females have been hypothesized to be extreme paedomorphic (i.e., larviform), however, females and larvae of the group are currently unknown. Here we provide the first association of adult male and immature life stages from the Puerto Rico using DNA barcoding, also collections-based associations and descriptions of immature Leptolycini and the first description of a paedomorphic female from the Virgin Islands. To carry out these life-stage associations we prepared an in-depth review of the Leptolycini fauna of the Puerto Rican bank (Puerto Rico and Virgin Islands). Several new taxa and taxonomic arrangements are proposed: Cessator crypticusnew species, Cessator tortolensisnew species, Cessator obrienorumnew species; Dracolycus chupacabranew genus and species, Dracolycus marshallinew species; Leptolycus falsoheterocornisnew species, and Leptolycus viensisnew species. Nanolycus gnomus Kazantsev is moved to Cessator gnomus (Kazantsev) new combination, rendering Nanolycus Kazantsev a new junior synonym of Cessator Kazantsev. The subgenus Baholycus Bocak is a new junior synonym of Leptolycus Leng and Mutchler. Leptolycus heterocornis var. flavicollis Leng and Mutchler is elevated to Leptolycus flavicollis Leng and Mutchler new status; Leptolycus (Leptolycus) albicauda Kazantsev is a new junior synonym of Leptolycus flavicollis Leng and Mutchler. An updated key to the adults and immature forms of Leptolycini from the Puerto Rican bank and a discussion on the importance of scientific collections in biodiversity studies is also provided.
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In this study, we describe a new Lampyridae subfamily, Chespiritoinae new subfamily, its sole genus Chespiritonew genus, and three new species: Chespirito zaragozainew species, Chespirito lloydinew species, and Chespirito ballantyneaenew species from the Neotropical portions of Mexico. Chespirito can be readily separated from all other known Lampyridae by the unique prosternum, characterized by being very wide, divided in the middle by a distinct suture forming two plates, with the anterior margin bearing a narrow inter-coxal process, the strongly setose filiform antennae, with antennomere III much smaller than all other antennomeres, the pronotum medially constricted (not in C. ballantyneae), with area adjacent to disc strongly punctate and with the presence of a strongly developed longitudinal carina. To test the placement of the genus within the Lampyridae, we obtained three genetic markers (18S rRNA, 28SrRNA, and cox1 mitochondrial DNA) from C. zaragozai and performed a maximum likelihood (ML) and Bayesian inference (BI) analysis. Our analyses rendered nearly identical tree topologies, with C. zaragozainew species recovered as an independent lineage as sister to Pollaclasis bifaria (Say) (Coleoptera, Lampyridae) + Cyphonocerus ruficollis Kiesenwetter (Coleoptera, Lampyridae)+Luciolinae, with a posterior probability of 96 for the BI analysis and UFBoot respectively of 91 for the ML analysis with the entire clade sister to Pterotus obscuripennis LeConte (Coleoptera, Lampyridae). A detailed examination of the morphology of Chespirito indicates that this lineage is divergent from all other known Lampyridae, which combined with the results of our analyses supports the erection of a new subfamily.
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Since the description of the genus Cheguevaria as incertae sedis (Lampyridae), the placement of these beetles has been uncertain. This study is the first to address the placement of this genus based on a molecular phylogenetic analysis. We used three genes (18S rRNA, rrnl mitochondrial DNA and cox1 mitochondrial DNA) and a maximum likelihood approach with W-IQ-TREE to support Cheguevaria as a member of the Lampyridae and recognize it as the sole genus in the new subfamily Cheguevariinae stat. nov.
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A new genus of Neotropical Lycidae, Currhaeus gen. nov., is herein proposed as the second Eurrhacini genus lacking parameres in male genitalia. Seven new species are described: Currhaeus striatus sp. nov., C. nigroapicalis sp. nov., C. championi sp. nov., C. tabascensis sp. nov., C. ruschii s . nov., C. polegattoi sp. nov., and C. paranaensis sp. nov. Illustrations of diagnostic characters and a key to species identification are presented. Parsimony and maximum likelihood analyses of morphological data demonstrated that Currhaeus gen. nov. belongs in the crown Eurrhacini. Implied weighting parsimony trees recovered Currhaeus as sister to Eurrhacus Waterhouse.
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The insect order Psocodea is a diverse lineage comprising both parasitic (Phthiraptera) and non-parasitic members (Psocoptera). The extreme age and ecological diversity of the group may be associated with major genomic changes, such as base compositional biases expected to affect phylogenetic inference. Divergent morphology between parasitic and non-parasitic members has also obscured the origins of parasitism within the order. We conducted a phylogenomic analysis on the order Psocodea utilizing both transcriptome and genome sequencing to obtain a data set of 2,370 orthologous genes. All phylogenomic analyses, including both concatenated and coalescent methods suggest a single origin of parasitism within the order Psocodea, resolving conflicting results from previous studies. This phylogeny allows us to propose a stable ordinal level classification scheme that retains significant taxonomic names present in historical scientific literature and reflects the evolution of the group as a whole. A dating analysis, with internal nodes calibrated by fossil evidence, suggests an origin of parasitism that predates the K-Pg boundary. Nucleotide compositional biases are detected in third and first codon positions and result in the anomalous placement of the Amphientometae as sister to Psocomorpha when all nucleotide sites are analyzed. Likelihood-mapping and quartet sampling methods demonstrate that base compositional biases can also have an effect on quartet-based methods.
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Despite the many substantial advances and progress in understanding the internal evolutionary relations of several Lycidae groups, the divergence time estimation of the family within Elateroidea remains subject of debate, with estimates to as early as the early to mid-Jurassic. Herein we describe Prototrichalus gen. nov., and three new species: Prototrichalus sepronai sp. nov., Prototrichalus meiyingae sp. nov.and Prototrichalus milleri sp. nov., from Burmese amber of the Cenomanian Age (mid-Cretaceous), supporting an age for the tribe of at least 100 Ma. The new taxa belong to the tribe Metriorrhynchini and are the oldest recognizable beetles from this lineage, indeed, among the oldest known representatives of the Lycidae. The discovery and description of Prototrichalus and its three species provides direct evidence of the group’s as a recognizable group in the mid-Cretaceous, much earlier than the recent predictions in the Late Cretaceous.
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The thoracic morphology of the troglobiontic leiodid species Troglocharinus ferreri (Cholevinae, Leptodirini) is described and documented in detail. The features are mainly discussed with respect to modifications linked with subterranean habits. Troglocharinus is assigned to the moderately modified pholeuonoid morphotype. The body is elongated and slender compared to epigean leiodids and also cave-dwelling species of Ptomaphagini. The legs are elongated, especially the hindlegs, though to a lesser degree than in the most advanced troglobiontic species. The prothorax is moderately elongated but otherwise largely unmodified. Its muscular system is strongly developed, with more muscle bundles that in free-living staphylinoid or hydrophiloid species. The pterothorax is greatly modified, especially the metathoracic flight apparatus. The meso- and metathoracic elements of the elytral locking device are well-developed, whereas the other notal parts are largely reduced. The mesonotum is simplified, with the triangular scutellar shield as the only distinctly developed part. The mesothoracic musculature is strongly reduced, with only 6 muscles compared to 12 or 13 in free-living staphylinoid or hydrophiloid species. The metanotum is greatly reduced, without a recognizable subdivision into prescutum scutum and scutellum. It is strongly narrowing laterally and lacks notal wing processes and other wing-related elements, but well-developed alacristae are present. The wings are reduced to small membranous flap-like structures inserted at the posterior end of the metanotum. A metapostnotum is not developed. Like in the case of the head, cave dwelling species of the related Ptomaphagini and Leptodirini show different trends of adaptations, with a compact ovoid or navicular body shape in the former, and a distinct trend towards elongation of the body and appendages in the latter tribe. Structural affinities of the thoraces of T. ferreri and the troglobiontic trechine carabid Sinaphaenops wangorum are mainly due to the reduced flight apparatus. The degree of muscle reduction in the pterothorax is very similar in both species.
Article
Cydistinae are a rare monogeneric beetle lineage from Asia with a convoluted history of classification, historically placed in various groups within the series Elateriformia. However, their position has never been rigorously tested. To resolve this long-standing puzzle, we are the first to present sequences of two nuclear and two mitochondrial markers for four species of Cydistinae to determine their phylogenetic position. We included these sequences in two rounds of analyses: one including a broad Elateriformia dataset to test placement at the superfamily/family level, and a second, including a richer, targeted sampling of presumed close relatives. Our results strongly support Cydistinae as sister to Phengodidae in a clade with Rhagophthalmidae. Based on our molecular phylogenetic results and examination of morphological characters, we hereby transfer the formerly unplaced Cydistinae into Phengodidae and provide diagnoses for the newly circumscribed Phengodidae, Cydistinae and Cydistus. Since both Phengodidae and Rhagophthalmidae have bioluminescent larvae and strongly neotenic females, similar features can be hypothesized for Cydistinae. Additionally, Cydistus minor is transferred to the new genus Microcydistus.
Article
The Neotropical genus Lycomorphon Pic, 1922 is revised. Eleven species are classified in the genus, six of them are redescribed, five species (Lycomorphon brasiliense sp. n., L. amazonicum sp. n., L. bimaculatum sp. n. L. bolivianum sp. n., L. fulvohumeralis sp. n.) are proposed as new to science. Lycomorphon elongaticolle v. diversicolle Pic, 1926 is raised to species status, and Idiopteron irregularis is transferred to the genus Lycomorphon. The new subgenus Spinolycus sg. n. is proposed within Lycomorphon.
Article
Drilini are soft-bodied predatory click-beetles (Elateridae: Agrypninae) with incompletely metamorphosed females. Due to divergent morphology, their classification has been contentious. We present the first densely sampled molecular phylogeny of Drilini based on nuclear and mitochondrial markers. Altogether, 44 species, representing all genera, were analysed using maximum likelihood and the Bayesian approach. Molecular analyses recovered five major clades that were also well supported by morphology. Afrotropical lineages mark deep splits. Most Palearctic species belong to a terminal clade. A few species of the predominantly Afrotropical Selasia are distributed from Arabia up to the Himalayas and Thailand. The origin of Drilini is dated to the Late Eocene (~35.5 Mya) and rapid radiation is identified from the Eocene/Oligocene to the Middle Miocene. We describe the gradual transformation of male morphological traits, e.g. the level of sclerotization, structure of mouthparts, loss of the thoracic interlocking mechanism, shortened elytra and expanded larviform abdomen. Five new genera, Austroselasia, Habeshaselasia, Illubaboria, Malacodrilus and Mashaselasia are proposed. Latoselasia, previously a subgenus of Wittmerselasia, is elevated to the genus rank. Five new species, Habeshaselasia nekemtensis, H. illubaborensis, Illubaboria bicolor, Malacodrilus hajeki and Mashaselasia aethiopica are described.
Article
Net-winged beetles (Coleoptera: Lycidae) are a diverse group of elateroids known for aposematism and neoteny. Phylogenetic analyses of morphological and molecular data have revealed different results with respect to within-group relationships. In this study, we recovered a highly supported phylogenomic phylogeny and identified seven subfamilies: Dexorinae stat.n., Calochrominae stat.n., Erotinae, Ateliinae, Lyci-nae, Lyropaeinae stat.n. and Metriorrhynchinae stat.n. Our results suggest that female neoteny evolved multiple times. Therefore, the development of similar morphological modifications in neotenics may be linked and may have produced characteristics such as body miniaturization, structural simplification, i.e. reduction of mouthparts, fewer antennomeres and palpomeres, uniquely shaped terminal palpomeres, shortened elytra, the loss of coadaptation between the elytra and pronotum, and others. Additional traits evolved in parallel due to similarities in biology, function and sexual selection. These characteristics include mimetic similarities, the presence of the rostrum, pronotal carinae and elytral costae, and the structure of male genitalia. By comparing the phylogenomic topology with the evolution of morphological characters, we were able to identify evolutionary trends in lycids and compare them with analogues for other neotenic elateroids. These traits have not been accepted as homoplasies due to the ambiguous phylogenetic signal from Sanger sequencing markers.
Article
We summarize knowledge of the weevil tribe Cleonini worldwide, including its monophyly, relationships, distribution, biology, immature stages, economic significance and paleontology. We score adult morphological characters for 79 of a total of 96 extant genus-group Cleonini taxa considered valid to date. The resulting matrix contains 121 parsimoniously informative characters scored for 145 ingroup (Cleonini) and 29 outgroup terminals. Maximum Parsimony (MP) and Bayesian Inference (BI) analyses consistently recover monophyletic Lixinae and Cleonini. Relationships within the latter remain unresolved with either 47 (BI) or 37 (MP) branches radiating from the tribe’s most recent common ancestor. Most of the speciose genera of Cleonini emerge as monophyletic in both BI and MP analyses (generic names followed by the number of terminals, then by BI posterior probability / MP bootstrap): Adosomus (5, 94/77), Asproparthenis (6, 99/98), Chromonotus (6, 98/85), Cleonis (3, 64/76), Coniocleonus (10, 95/41), Conorhynchus (5, 95/51), Cyphoclenus (4, 65/76), Maximus (4, 84/68), Mecaspis (4, 95/91), Scaphomorphus (4, 90/84), Temnorhinus (8, 99/62) and Xanthochelus (6, 84/71). The genera Pseudocleonus (6, -/26) and Stephanocleonus (22, -/23) are not recovered in BI and weakly supported in MP. No genera are here added to, or removed from, Cleonini. We suggest that adult morphology of Cleonini was subject to widespread homoplasy obscuring the phylogenetic signal of morphological characters. Unlike the rest of Lixinae, all extant Cleonini are hypothesised to be flightless, even though often being macropterous. All 145 ingroup terminals are illustrated in three standard views; images of the type species of 15 of the 17 genus-group taxa that are not represented in our analysis are provided.
Article
Model-based molecular phylogenetics plays an important role in comparisons of genomic data, and model selection is a key step in all such analyses. We present ModelFinder, a fast model-selection method that greatly improves the accuracy of phylogenetic estimates by incorporating a model of rate heterogeneity across sites not previously considered in this context and by allowing concurrent searches of model space and tree space.