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Mastigocoleidae fam. nov., a New Mesozoic Beetle Family and the Early Evolution of Dryopoidea (Coleoptera)
Abstract and Figures
With some 3,700 described species, Dryopoidea are a moderately diverse superfamily of beetles whose position within basal Polyphaga has been historically difficult to elucidate. Members of most extant dryopoid families are set apart from the majority of other polyphagans by their association with aquatic habitats, but little is known about the origin of these derived life habits and the phylogeny of the superfamily. Here we describe Mastigocoleidae Tihelka, Jäch, Kundrata & Cai fam. nov., a new family of Mesozoic dryopoids represented by fossils from the Cretaceous Yixian Formation in northeastern China (undescribed species; ~125 Ma), Crato Formation in northeastern Brazil (Mastigocoleus rhinoceros Tihelka & Cai gen. et sp. nov.; ~113 Ma), and amber from northern Myanmar (Mastigocoleus resinicola Tihelka & Cai gen. et sp. nov. and Cretaceocoleus saetosus Tihelka, Kundrata & Cai gen. et sp. nov.; ~99 Ma). Integrating the findings of recent molecular and morphological phylogenetic analyses, we recover Mastigocoleidae as an early-diverging dryopoid clade sister to the families Lutrochidae and Dryopidae, or less likely as a group of putative stem-dryopoids. Mastigocoleidae are most distinctly separated from all other dryopoid families by their whip-like antennae, with 11 antennomeres, reaching to the pronotal base, and with the scape broadest and longest, a short pedicel, and antennomeres II–XI more or less distinctively gradually tapering toward the apex. Mastigocoleidae indicate that the last common ancestor of Dryopoidea was likely terrestrial in the adult stage, and document character acquisitions associated with a specialization for aquatic life.
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... The family Ptilodactylidae is a small group of beetles, with most representatives living near water, often with larvae that have special adaptations for survival in aquatic habitats [1,2]. Ptilodactylidae belong to the superfamily Dryopoidea [3], but the detailed relationships of the family remain unclear [4][5][6]. The subfamilial and generic classification of Ptilodactylidae is in urgent need of revision [1, [7][8][9]. ...
... One of the unanswered questions regarding A. monachus is its systematic placement. Although it is currently widely accepted as a member of the family Ptilodactylidae, its relationships remain unclear due to the unique larval morphology, lack of a stable phylogenetic hypothesis for Dryopidea, and widely questioned monophyly of the family Ptilodactylidae [4][5][6]20,23,28]. Lawrence and Newton [26] (p. ...
... They placed Araeopidius in a clade with Cladotominae, Chelonariidae, Dryopidae, Psephenidae, Lutrochidae, and Elmidae (sharing an abdominal operculum) [28], in various clades including various elateriform groups [23], or in clades consisting of a part of Ptilodactylidae, or some Ptilodactylidae and Chelonariidae [20]. Although A. monachus has been sequenced in a barcoding project on Alaskan non-marine arthropods [67] and in an unpublished barcoding project at the Canadian National Collection (Appendix A), it has not been included in any molecular phylogenetic or phylogenomic study so far, e.g., [3,4,6,21,68]. ...
Araeopidius monachus (LeConte, 1874) is the only species in the subfamily Araeopidiinae within the beetle family Ptilodactylidae. Its geographic distribution is mainly along the western coast of North America, where adults are uncommon. A diagnosis is provided along with detailed collection records highlighting its seasonality, elevational range, plant associations, and collection methods. Collection records from larvae and adults indicate a three-year life cycle. Digestive tract dissections show that the larvae consume woody material while the adults probably do not feed. Additionally, we briefly discuss the problems and prospects for research of this enigmatic species.
... A considerable number of Cretaceous fossils of Dryopoidea were described recently, for instance, including a new genus of Ptilodactylidae (Chatzimanolis, Cashion, et al., 2012), species of Limnichidae (Li, Ruta, et al., 2022;Yu et al., 2018), Psephenidae Li, Yu, et al., 2022), Elmidae (Cai, Maier, & Huang, 2018) and Heteroceridae (Li, Tihelka, et al., 2020). Tihelka et al. (2022) described a new family of dryopoid beetles, Mastigocoleidae, based on compression fossils and amber inclusions from three Early to Late Cretaceous deposits. ...
Recent progress in beetle palaeontology has incited us to re-address the evolutionary history of the group. The Permian †Tshekardocoleidae had elytra that covered the posterior body in a loose tent-like manner. The formation of elytral epipleura and a tight fit of elytra and abdomen were important evolutionary transformations in the Middle Permian, resulting in a tightly enclosed subelytral space. Permian families were likely associated with dead wood of gymnospermous trees. The end-Permian extinction event resulted in a turnover in the composition of beetle faunas, especially a decline of large-bodied wood-associated forms. Adephaga and Myxophaga underwent a first wave of diversification in the Triassic. Polyphaga are very rare in this period. The first wave of diversification of this suborder occurs in the Jurassic, with fossils of Elateriformia, Staphyliniformia and Cucujiformia. The Cretaceous fossil record has been tremendously enriched by the discovery of amber inclusions. Numerous fossils represent all major polyphagan lineages and also the remaining suborders. Improved analytical methods for documenting and placing extinct taxa are discussed. Different factors have played a role in the diversification of beetles. The enormous number of species associated with flowering plants, and timing and patterns of diversification in phytophagous lineages indicate that the angio-sperm radiation played a major role in beetle macroevolution. Moreover, the evolution of intimate partnerships with symbionts and the acquisition of novel genes-obtained from fungi and bacteria via horizontal gene transfers-facilitated the use of plant material as a food source and were key innovations in the diversification of plant-feeding beetles.
... The subfamily Dryopoidea includes the following families: Chelonariidae, Dryopidae, Elmidae Curtis, 1840 [33], Protelmidae Jeannel, 1950 [34], Eulichadidae Crowson, 1973 [35], Heteroceridae MacLeay, 1825 [36], Limnichidae, Lutrochidae, Psephenidae Lacordaire, 1854 [37], Ptilodactylidae Laporte, 1836 [38], Callirhipidae Emden, 1924 [39], Cneoglossidae Champion, 1897 [40] and †Mastigocoleidae Tihelka, Jäch, Kundrata et Cai, 2022 [41]. This superfamily was grounded and divided by Crowson into two groups of families (dryopid and elmid groups) and frequently with some differences in composition, argumentation [42,43], etc. ...
Burmochares groehnigen. et sp. nov., the oldest known representative of the subfamily Limnichinae and tribe Limnichini, is described from the Cretaceous amber of northern Myanmar. The new genus Hernandocharesgen. nov. is proposed for Platypelochares electricus Hernando, Szawaryn et Ribera, 2018 (type species of this new genus) from the Eocene Baltic amber. The structurally similar genera Platypelochares,Burmocharesgen.nov., Hernandocharesgen. nov., some other limnichines (Polyphaga) and some torridincollids (Myxophaga) are thought to be connected by their preference for similar habitats and lifestyle.
In order to place newly discovered fossil taxa ( Palaeosymbius gen. nov. with P. groehni and P. mesozoicus spp. nov.) from the mid‐Cretaceous amber from northern Myanmar, we investigated the relations of extant and extinct lineages of the coccinellid group of Coccinelloidea with emphasis on the family Anamorphidae. We assembled a taxonomic sampling of 34 taxa, including 15 genera and 19 species of Anamorphidae, the most comprehensive sampling of Anamorphidae at the generic level in a phylogenetic analysis. A morphological dataset of 47 characters was built as well as a molecular alignment of 7140 bp including fragments of eight genes ( 12S , 16S , 18S , 28S , COI , COII , H3 and CAD ). Five anamorphid and one endomychid species were sequenced for the first time and added to the dataset. We performed parsimony‐based analysis of the morphological dataset and Bayesian inference analysis of the combined matrix (morphological plus molecular data). Our results confirm that Palaeosymbius belongs to Anamorphidae and represents the oldest known member of this family so far. Among Anamorphidae, Symbiotes (with extant and known Eocene species) was recovered as the most probable closest relative of Palaeosymbius. Our morphological studies additionally revealed the presence of probable glandular openings in the anterolateral corners of the pronotal margins in Asymbius sp. and Anamorphus sp., representing the first report of secretory openings in the family Anamorphidae. Similar openings are found in other cucujiform beetles such as Cryptophagidae and Boganiidae with possible defensive purposes.
More than 4700 nominal family-group names (including names for fossils and ichnotaxa) are nomenclaturally available in the order Coleoptera. Since each family-group name is based on the concept of its type genus, we argue that the stability of names used for the classification of beetles depends on accurate nomenclatural data for each type genus. Following a review of taxonomic literature, with a focus on works that potentially contain type species designations, we provide a synthesis of nomenclatural data associated with the type genus of each nomenclaturally available family-group name in Coleoptera. For each type genus the author(s), year of publication, and page number are given as well as its current status (i.e., whether treated as valid or not) and current classification.
Information about the type species of each type genus and the type species fixation (i.e., fixed originally or subsequently, and if subsequently, by whom) is also given. The original spelling of the family-group name that is based on each type genus is included, with its author(s), year, and stem. We append a list of nomenclaturally available family-group names presented in a classification scheme. Because of the importance of the Principle of Priority in zoological nomenclature, we provide information on the date of publication of the references cited in this work, when known. Several nomenclatural issues emerged during the course of this work. We therefore appeal to the community of coleopterists to submit applications to the International Commission on Zoological Nomenclature (henceforth “Commission”) in order to permanently resolve some of the problems outlined here.
The following changes of authorship for type genera are implemented here (these changes do not affect the concept of each type genus): CHRYSOMELIDAE: Fulcidax Crotch, 1870 (previously credited to “Clavareau, 1913”); CICINDELIDAE: Euprosopus W.S. MacLeay, 1825 (previously credited to “Dejean, 1825”); COCCINELLIDAE: Alesia Reiche, 1848 (previously credited to “Mulsant, 1850”); CURCULIONIDAE: Arachnopus Boisduval, 1835 (previously credited to “Guérin-Méneville, 1838”); ELATERIDAE: Thylacosternus Gemminger, 1869 (previously credited to “Bonvouloir, 1871”); EUCNEMIDAE: Arrhipis Gemminger, 1869 (previously credited to “Bonvouloir, 1871”), Mesogenus Gemminger, 1869 (previously credited to “Bonvouloir, 1871”); LUCANIDAE: Sinodendron Hellwig, 1791 (previously credited to “Hellwig, 1792”); PASSALIDAE: Neleides Harold, 1868 (previously credited to “Kaup, 1869”), Neleus Harold, 1868 (previously credited to “Kaup, 1869”), Pertinax Harold, 1868 (previously credited to “Kaup, 1869”), Petrejus Harold, 1868 (previously credited to “Kaup, 1869”), Undulifer Harold, 1868 (previously credited to “Kaup, 1869”), Vatinius Harold, 1868 (previously credited to “Kaup, 1869”); PTINIDAE: Mezium Leach, 1819 (previously credited to “Curtis, 1828”); PYROCHROIDAE: Agnathus Germar, 1818 (previously credited to “Germar, 1825”); SCARABAEIDAE: Eucranium Dejean, 1833 (previously “Brullé, 1838”).
The following changes of type species were implemented following the discovery of older type species fixations (these changes do not pose a threat to nomenclatural stability): BOLBOCERATIDAE: Bolbocerus bocchus Erichson, 1841 for Bolbelasmus Boucomont, 1911 (previously Bolboceras gallicum Mulsant, 1842); BUPRESTIDAE: Stigmodera guerinii Hope, 1843 for Neocuris Saunders, 1868 (previously Anthaxia fortnumi Hope, 1846), Stigmodera peroni Laporte & Gory, 1837 for Curis Laporte & Gory, 1837 (previously Buprestis caloptera Boisduval, 1835); CARABIDAE: Carabus elatus Fabricius, 1801 for Molops Bonelli, 1810 (previously Carabus terricola Herbst, 1784 sensu Fabricius, 1792); CERAMBYCIDAE: Prionus palmatus Fabricius, 1792 for Macrotoma Audinet-Serville, 1832 (previously Prionus serripes Fabricius, 1781); CHRYSOMELIDAE: Donacia equiseti Fabricius, 1798 for Haemonia Dejean, 1821 (previously Donacia zosterae Fabricius, 1801), Eumolpus ruber Latreille, 1807 for Euryope Dalman, 1824 (previously Cryptocephalus rubrifrons Fabricius, 1787), Galeruca affinis Paykull, 1799 for Psylliodes Latreille, 1829 (previously Chrysomela chrysocephala Linnaeus, 1758); COCCINELLIDAE: Dermestes rufus Herbst, 1783 for Coccidula Kugelann, 1798 (previously Chrysomela scutellata Herbst, 1783); CRYPTOPHAGIDAE: Ips caricis G.-A. Olivier, 1790 for Telmatophilus Heer, 1841 (previously Cryptophagus typhae Fallén, 1802), Silpha evanescens Marsham, 1802 for Atomaria Stephens, 1829 (previously Dermestes nigripennis Paykull, 1798); CURCULIONIDAE: Bostrichus cinereus Herbst, 1794 for Crypturgus Erichson, 1836 (previously Bostrichus pusillus Gyllenhal, 1813); DERMESTIDAE: Dermestes trifasciatus Fabricius, 1787 for Attagenus Latreille, 1802 (previously Dermestes pellio Linnaeus, 1758); ELATERIDAE: Elater sulcatus Fabricius, 1777 for Chalcolepidius Eschscholtz, 1829 (previously Chalcolepidius zonatus Eschscholtz, 1829); ENDOMYCHIDAE: Endomychus rufitarsis Chevrolat, 1835 for Epipocus Chevrolat, 1836 (previously Endomychus tibialis Guérin-Méneville, 1834); EROTYLIDAE: Ips humeralis Fabricius, 1787 for Dacne Latreille, 1797 (previously Dermestes bipustulatus Thunberg, 1781); EUCNEMIDAE: Fornax austrocaledonicus Perroud & Montrouzier, 1865 for Mesogenus Gemminger, 1869 (previously Mesogenus mellyi Bonvouloir, 1871); GLAPHYRIDAE: Melolontha serratulae Fabricius, 1792 for Glaphyrus Latreille, 1802 (previously Scarabaeus maurus Linnaeus, 1758); HISTERIDAE: Hister striatus Forster, 1771 for Onthophilus Leach, 1817 (previously Hister sulcatus Moll, 1784); LAMPYRIDAE: Ototreta fornicata E. Olivier, 1900 for Ototreta E. Olivier, 1900 (previously Ototreta weyersi E. Olivier, 1900); LUCANIDAE: Lucanus cancroides Fabricius, 1787 for Lissotes Westwood, 1855 (previously Lissotes menalcas Westwood, 1855); MELANDRYIDAE: Nothus clavipes G.-A. Olivier, 1812 for Nothus G.-A. Olivier, 1812 (previously Nothus praeustus G.-A. Olivier, 1812); MELYRIDAE: Lagria ater Fabricius, 1787 for Enicopus Stephens, 1830 (previously Dermestes hirtus Linnaeus, 1767); NITIDULIDAE: Sphaeridium luteum Fabricius, 1787 for Cychramus Kugelann, 1794 (previously Strongylus quadripunctatus Herbst, 1792); OEDEMERIDAE: Helops laevis Fabricius, 1787 for Ditylus Fischer, 1817 (previously Ditylus helopioides Fischer, 1817 [sic]); PHALACRIDAE: Sphaeridium aeneum Fabricius, 1792 for Olibrus Erichson, 1845 (previously Sphaeridium bicolor Fabricius, 1792); RHIPICERIDAE: Sandalus niger Knoch, 1801 for Sandalus Knoch, 1801 (previously Sandalus petrophya Knoch, 1801); SCARABAEIDAE: Cetonia clathrata G.-A. Olivier, 1792 for Inca Lepeletier & Audinet-Serville, 1828 (previously Cetonia ynca Weber, 1801); Gnathocera vitticollis W. Kirby, 1825 for Gnathocera W. Kirby, 1825 (previously Gnathocera immaculata W. Kirby, 1825); Melolontha villosula Illiger, 1803 for Chasmatopterus Dejean, 1821 (previously Melolontha hirtula Illiger, 1803); STAPHYLINIDAE: Staphylinus politus Linnaeus, 1758 for Philonthus Stephens, 1829 (previously Staphylinus splendens Fabricius, 1792); ZOPHERIDAE: Hispa mutica Linnaeus, 1767 for Orthocerus Latreille, 1797 (previously Tenebrio hirticornis DeGeer, 1775).
The discovery of type species fixations that are older than those currently accepted pose a threat to nomenclatural stability (an application to the Commission is necessary to address each problem): CANTHARIDAE: Malthinus Latreille, 1805, Malthodes Kiesenwetter, 1852; CARABIDAE: Bradycellus Erichson, 1837, Chlaenius Bonelli, 1810, Harpalus Latreille, 1802, Lebia Latreille, 1802, Pheropsophus Solier, 1834, Trechus Clairville, 1806; CERAMBYCIDAE: Callichroma Latreille, 1816, Callidium Fabricius, 1775, Cerasphorus Audinet-Serville, 1834, Dorcadion Dalman, 1817, Leptura Linnaeus, 1758, Mesosa Latreille, 1829, Plectromerus Haldeman, 1847; CHRYSOMELIDAE: Amblycerus Thunberg, 1815, Chaetocnema Stephens, 1831, Chlamys Knoch, 1801, Monomacra Chevrolat, 1836, Phratora Chevrolat, 1836, Stylosomus Suffrian, 1847; COLONIDAE: Colon Herbst, 1797; CURCULIONIDAE: Cryphalus Erichson, 1836, Lepyrus Germar, 1817; ELATERIDAE: Adelocera Latreille, 1829, Beliophorus Eschscholtz, 1829; ENDOMYCHIDAE: Amphisternus Germar, 1843, Dapsa Latreille, 1829; GLAPHYRIDAE: Anthypna Eschscholtz, 1818; HISTERIDAE: Hololepta Paykull, 1811, Trypanaeus Eschscholtz, 1829; LEIODIDAE: Anisotoma Panzer, 1796, Camiarus Sharp, 1878, Choleva Latreille, 1797; LYCIDAE: Calopteron Laporte, 1838, Dictyoptera Latreille, 1829; MELOIDAE: Epicauta Dejean, 1834; NITIDULIDAE: Strongylus Herbst, 1792; SCARABAEIDAE: Anisoplia Schönherr, 1817, Anticheira Eschscholtz, 1818, Cyclocephala Dejean, 1821, Glycyphana Burmeister, 1842, Omaloplia Schönherr, 1817, Oniticellus Dejean, 1821, Parachilia Burmeister, 1842, Xylotrupes Hope, 1837; STAPHYLINIDAE: Batrisus Aubé, 1833, Phloeonomus Heer, 1840, Silpha Linnaeus, 1758; TENEBRIONIDAE: Bolitophagus Illiger, 1798, Mycetochara Guérin-Méneville, 1827.
Type species are fixed for the following nominal genera: ANTHRIBIDAE: Decataphanes gracilis Labram & Imhoff, 1840 for Decataphanes Labram & Imhoff, 1840; CARABIDAE: Feronia erratica Dejean, 1828 for Loxandrus J.L. LeConte, 1853; CERAMBYCIDAE: Tmesisternus oblongus Boisduval, 1835 for Icthyosoma Boisduval, 1835; CHRYSOMELIDAE: Brachydactyla annulipes Pic, 1913 for Pseudocrioceris Pic, 1916, Cassida viridis Linnaeus, 1758 for Evaspistes Gistel, 1856, Ocnoscelis cyanoptera Erichson, 1847 for Ocnoscelis Erichson, 1847, Promecotheca petelii Guérin-Méneville, 1840 for Promecotheca Guérin- Méneville, 1840; CLERIDAE: Attelabus mollis Linnaeus, 1758 for Dendroplanetes Gistel, 1856; CORYLOPHIDAE: Corylophus marginicollis J.L. LeConte, 1852 for Corylophodes A. Matthews, 1885; CURCULIONIDAE: Hoplorhinus melanocephalus Chevrolat, 1878 for Hoplorhinus Chevrolat, 1878; Sonnetius binarius Casey, 1922 for Sonnetius Casey, 1922; ELATERIDAE: Pyrophorus melanoxanthus Candèze, 1865 for Alampes Champion, 1896; PHYCOSECIDAE: Phycosecis litoralis Pascoe, 1875 for Phycosecis Pascoe, 1875; PTILODACTYLIDAE: Aploglossa sallei Guérin-Méneville, 1849 for Aploglossa Guérin-Méneville, 1849, Colobodera ovata Klug, 1837 for Colobodera Klug, 1837; PTINIDAE: Dryophilus anobioides Chevrolat, 1832 for Dryobia Gistel, 1856; SCARABAEIDAE: Achloa helvola Erichson, 1840 for Achloa Erichson, 1840, Camenta obesa Burmeister, 1855 for Camenta Erichson, 1847, Pinotus talaus Erichson, 1847 for Pinotus Erichson, 1847, Psilonychus ecklonii Burmeister, 1855 for Psilonychus Burmeister, 1855.
New replacement name: CERAMBYCIDAE: Basorus Bouchard & Bousquet, nom. nov. for Sobarus Harold, 1879.
New status: CARABIDAE: KRYZHANOVSKIANINI Deuve, 2020, stat. nov. is given the rank of tribe instead of subfamily since our classification uses the rank of subfamily for PAUSSINAE rather than family rank; CERAMBYCIDAE: Amymoma Pascoe, 1866, stat. nov. is used as valid over Neoamymoma Marinoni, 1977, Holopterus Blanchard, 1851, stat. nov. is used as valid over Proholopterus Monné, 2012; CURCULIONIDAE: Phytophilus Schönherr, 1835, stat. nov. is used as valid over the unnecessary new replacement name Synophthalmus Lacordaire, 1863; EUCNEMIDAE: Nematodinus Lea, 1919, stat. nov. is used as valid instead of Arrhipis Gemminger, 1869, which is a junior homonym.
Details regarding additional nomenclatural issues that still need to be resolved are included in the entry for each of these type genera: BOSTRICHIDAE: Lyctus Fabricius, 1792; BRENTIDAE: Trachelizus Dejean, 1834; BUPRESTIDAE: Pristiptera Dejean, 1833; CANTHARIDAE: Chauliognathus Hentz, 1830, Telephorus Schäffer, 1766; CARABIDAE: Calathus Bonelli, 1810, Cosnania Dejean, 1821, Dicrochile Guérin-Méneville, 1847, Epactius D.H. Schneider, 1791, Merismoderus Westwood, 1847, Polyhirma Chaudoir, 1850, Solenogenys Westwood, 1860, Zabrus Clairville, 1806; CERAMBYCIDAE: Ancita J. Thomson, 1864, Compsocerus Audinet-Serville, 1834, Dorcadodium Gistel, 1856, Glenea Newman, 1842; Hesperophanes Dejean, 1835, Neoclytus J. Thomson, 1860, Phymasterna Laporte, 1840, Tetrops Stephens, 1829, Zygocera Erichson, 1842; CHRYSOMELIDAE: Acanthoscelides Schilsky, 1905, Corynodes Hope, 1841, Edusella Chapuis, 1874; Hemisphaerota Chevrolat, 1836; Physonota Boheman, 1854, Porphyraspis Hope, 1841; CLERIDAE: Dermestoides Schäffer, 1777; COCCINELLIDAE: Hippodamia Chevrolat, 1836, Myzia Mulsant, 1846, Platynaspis L. Redtenbacher, 1843; CURCULIONIDAE: Coeliodes Schönherr, 1837, Cryptoderma Ritsema, 1885, Deporaus Leach, 1819, Epistrophus Kirsch, 1869, Geonemus Schönherr, 1833, Hylastes Erichson, 1836; DYTISCIDAE: Deronectes Sharp, 1882, Platynectes Régimbart, 1879; EUCNEMIDAE: Dirhagus Latreille, 1834; HYBOSORIDAE: Ceratocanthus A. White, 1842; HYDROPHILIDAE: Cyclonotum Erichson, 1837; LAMPYRIDAE: Luciola Laporte, 1833; LEIODIDAE: Ptomaphagus Hellwig, 1795; LUCANIDAE: Leptinopterus Hope, 1838; LYCIDAE: Cladophorus Guérin-Méneville, 1830, Mimolibnetis Kazantsev, 2000; MELOIDAE: Mylabris Fabricius, 1775; NITIDULIDAE: Meligethes Stephens, 1829; PTILODACTYLIDAE: Daemon Laporte, 1838; SCARABAEIDAE: Allidiostoma Arrow, 1940, Heterochelus Burmeister, 1844, Liatongus Reitter, 1892, Lomaptera Gory & Percheron, 1833, Megaceras Hope, 1837, Stenotarsia Burmeister, 1842; STAPHYLINIDAE: Actocharis Fauvel, 1871, Aleochara Gravenhorst, 1802; STENOTRACHELIDAE: Stenotrachelus Berthold, 1827; TENEBRIONIDAE: Cryptochile Latreille, 1828, Heliopates Dejean, 1834, Helops Fabricius, 1775.
First Reviser actions deciding the correct original spelling: CARABIDAE: Aristochroodes Marcilhac, 1993 (not Aritochroodes ); CERAMBYCIDAE: Dorcadodium Gistel, 1856 (not Dorcadodion ), EVODININI Zamoroka, 2022 (not EVODINIINI); CHRYSOMELIDAE: Caryopemon Jekel, 1855 (not Carpopemon ), Decarthrocera Laboissière, 1937 (not Decarthrocerina ); CICINDELIDAE: Odontocheila Laporte, 1834 (not Odontacheila ); CLERIDAE: CORMODINA Bartlett, 2021 (not CORMODIINA), Orthopleura Spinola, 1845 (not Orthoplevra , not Orthopleuva ); CURCULIONIDAE: Arachnobas Boisduval, 1835 (not Arachnopus ), Palaeocryptorhynchus Poinar, 2009 (not Palaeocryptorhynus ); DYTISCIDAE: Ambarticus Yang et al., 2019 and AMBARTICINI Yang et al., 2019 (not Ambraticus , not AMBRATICINI); LAMPYRIDAE: Megalophthalmus G.R. Gray, 1831 (not Megolophthalmus , not Megalopthalmus ); SCARABAEIDAE: Mentophilus Laporte, 1840 (not Mintophilus , not Minthophilus ), Pseudadoretus dilutellus Semenov, 1889 (not P. ditutellus ).
While the correct identification of the type species is assumed, in some cases evidence suggests that species were misidentified when they were fixed as the type of a particular nominal genus. Following the requirements of Article 70.3.2 of the International Code of Zoological Nomenclature we hereby fix the following type species (which in each case is the taxonomic species actually involved in the misidentification): ATTELABIDAE: Rhynchites cavifrons Gyllenhal, 1833 for Lasiorhynchites Jekel, 1860; BOSTRICHIDAE: Ligniperda terebrans Pallas, 1772 for Apate Fabricius, 1775; BRENTIDAE: Ceocephalus appendiculatus Boheman, 1833 for Uroptera Berthold, 1827; BUPRESTIDAE: Buprestis undecimmaculata Herbst, 1784 for Ptosima Dejean, 1833; CARABIDAE: Amara lunicollis Schiødte, 1837 for Amara Bonelli, 1810, Buprestis connexus Geoffroy, 1785 for Polistichus Bonelli, 1810, Carabus atrorufus Strøm, 1768 for Patrobus Dejean, 1821, Carabus gigas Creutzer, 1799 for Procerus Dejean, 1821, Carabus teutonus Schrank, 1781 for Stenolophus Dejean, 1821, Carenum bonellii Westwood, 1842 for Carenum Bonelli, 1813, Scarites picipes G.-A. Olivier, 1795 for Acinopus Dejean, 1821, Trigonotoma indica Brullé, 1834 for Trigonotoma Dejean, 1828; CERAMBYCIDAE: Cerambyx lusitanus Linnaeus, 1767 for Exocentrus Dejean, 1835, Clytus supernotatus Say, 1824 for Psenocerus J.L. LeConte, 1852; CICINDELIDAE: Ctenostoma jekelii Chevrolat, 1858 for Ctenostoma Klug, 1821; CURCULIONIDAE: Cnemogonus lecontei Dietz, 1896 for Cnemogonus J.L. LeConte, 1876; Phloeophagus turbatus Schönherr, 1845 for Phloeophagus Schönherr, 1838; GEOTRUPIDAE: Lucanus apterus Laxmann, 1770 for Lethrus Scopoli, 1777; HISTERIDAE: Hister rugiceps Duftschmid, 1805 for Hypocaccus C.G. Thomson, 1867; HYBOSORIDAE: Hybosorus illigeri Reiche, 1853 for Hybosorus W.S. MacLeay, 1819; HYDROPHILIDAE: Hydrophilus melanocephalus G.-A. Olivier, 1793 for Enochrus C.G. Thomson, 1859; MYCETAEIDAE: Dermestes subterraneus Fabricius, 1801 for Mycetaea Stephens, 1829; SCARABAEIDAE: Aulacium carinatum Reiche, 1841 for Mentophilus Laporte, 1840, Phanaeus vindex W.S. MacLeay, 1819 for Phanaeus W.S. MacLeay, 1819, Ptinus germanus Linnaeus, 1767 for Rhyssemus Mulsant, 1842, Scarabaeus latipes Guérin-Méneville, 1838 for Cheiroplatys Hope, 1837; STAPHYLINIDAE: Scydmaenus tarsatus P.W.J. Müller & Kunze, 1822 for Scydmaenus Latreille, 1802.
New synonyms: CERAMBYCIDAE: CARILIINI Zamoroka, 2022, syn. nov. of ACMAEOPINI Della Beffa, 1915, DOLOCERINI Özdikmen, 2016, syn. nov. of BRACHYPTEROMINI Sama, 2008, PELOSSINI Tavakilian, 2013, syn. nov. of LYGRINI Sama, 2008, PROHOLOPTERINI Monné, 2012, syn. nov. of HOLOPTERINI Lacordaire, 1868.
This is a supplement to the Burmese (Myanmar) amber checklist and bibliography covering taxa described or recorded during 2022, plus a couple of earlier records that were missed previously. Up to the end of 2022, 2,524 species have been recorded from Kachin amber, of which 350 were named in 2022; ten species have been recorded from older Hkamti amber, of which two were named in 2022 (one species known from both Hkamti and Kachin amber). Another 17 species were named in 2022 though it is uncertain whether they are in Kachin or Hkamti amber. In total 368 species were named from Cretaceous amber from Myanmar in 2022.
Beetles constitute the most biodiverse animal order with over 380 000 described species and possibly several million more yet unnamed. Recent phylogenomic studies have arrived at considerably incongruent topologies and widely varying estimates of divergence dates for major beetle clades. Here, we use a dataset of 68 single-copy nuclear protein-coding (NPC) genes sampling 129 out of the 193 recognized extant families as well as the first comprehensive set of fully justified fossil calibrations to recover a refined timescale of beetle evolution. Using phylogenetic methods that counter the effects of compositional and rate heterogeneity, we recover a topology congruent with morphological studies, which we use, combined with other recent phylogenomic studies, to propose several formal changes in the classification of Coleoptera: Scirtiformia and Scirtoidea sensu nov., Clambiformia ser. nov. and Clamboidea sensu nov., Rhinorhipiformia ser. nov., Byrrhoidea sensu nov., Dryopoidea stat. res., Nosodendriformia ser. nov. and Staphyliniformia sensu nov., and Erotyloidea stat. nov., Nitiduloidea stat. nov. and Cucujoidea sensu nov., alongside changes below the superfamily level. Our divergence time analyses recovered a late Carboniferous origin of Coleoptera, a late Palaeozoic origin of all modern beetle suborders and a Triassic–Jurassic origin of most extant families, while fundamental divergences within beetle phylogeny did not coincide with the hypothesis of a Cretaceous Terrestrial Revolution.
Bioluminescent beetles of the superfamily Elateroidea (fireflies, fire beetles, glow-worms) are the most speciose group of terrestrial light-producing animals. The evolution of bioluminescence in elateroids is associated with unusual morphological modifications, such as soft-bodiedness and neoteny, but the fragmentary nature of the fossil record discloses little about the origin of these adaptations. We report the discovery of a new bioluminescent elateroid beetle family from the mid-Cretaceous of northern Myanmar (ca 99 Ma), Cretophengodidae fam. nov. Cretophengodes azari gen. et sp. nov. belongs to the bioluminescent lampyroid clade, and would appear to represent a transitional fossil linking the soft-bodied Phengodidae + Rhagophthalmidae clade and hard-bodied elateroids. The fossil male possesses a light organ on the abdomen which presumably served a defensive function, documenting a Cretaceous radiation of bioluminescent beetles coinciding with the diversification of major insectivore groups such as frogs and stem-group birds. The discovery adds a key branch to the elateroid tree of life and sheds light on the evolution of soft-bodiedness and the historical biogeography of elateroid beetles.
Significance
Competition for common resources can make some species groups thrive and others decline. Flowering plants rose to dominance between 125 and 80 Ma, undergoing an explosive radiation that is believed to have impacted long-established plant groups like gymnosperms. Here, we show that the decline of conifers is strongly and directly linked to the increasing diversity of flowering plants. Both the fossil record and molecular data converge in clarifying the effects of abiotic or biotic factors on the speciation and extinction rates of conifers. These results imply that long-term biological interactions through clade competition can play a more important role in the rise and demise of major organism groups than mass extinctions.
The monospecific family Mysteriomorphidae was recently described based on two fossil specimens from the Late Cretaceous Kachin amber of northern Myanmar. The family was placed in Elateriformia incertae sedis without a clear list of characters that define it either in Elateroidea or in Byrrhoidea. We report here four additional adult specimens of the same lineage, one of which was described using a successful reconstruction from a CT-scan analysis to better observe some characters. The new specimens enabled us to considerably improve the diagnosis of Mysteriomorphidae. The family is definitively placed in Elateroidea, and we hypothesize its close relationship with Elateridae. Similarly, there are other fossil families of beetles that are exclusively described from Cretaceous ambers. These lineages may have been evolutionarily replaced by the ecological revolution launched by angiosperms that introduced new co-associations with taxa. These data indicate a macroevolutionary pattern of replacement that could be extended to other insect groups.
A new wood wasp, Cratosirex sennlaubi gen. et sp. nov., is described and figured from one specimen collected from the Lower Cretaceous Crato Formation in northeastern Brazil. This new genus is placed in the new siricid subfamily Cratosiricinae subfam. nov., based on a combination of plesiomorphic and autapomorphic characters. The presence of small and sub-equal forewing cells 1R1 and 2R1 is a synapomorphy with the extant subfamily Siricinae, absent in the other extinct subfamilies †Auliscinae and †Gigasiricinae, supporting a sister group relationships with the Siricinae. Our new discovery expands the distribution range of Siricidae fossil records, highlights the antiquity of the family, and emphasizes the need for more studies of this particular insect lineage in the Mesozoic deposits. Currently, all the representatives of the crown group of the extant Siricidae are Cenozoic.
New Zealand is an island continent that completed its split from the Gondwanan continent at 52 Ma, harbouring an iconic biota of tuatara, kiwi and weta. The sooty mould community is a distinctive trophic element of New Zealand forest ecosystems that is driven by plant-feeding sternorrhynchan Hemiptera. These produce honeydew, which supports fungal growth, which in turn supports numerous endemic invertebrates, including endemic New Zealand beetle families. Ancient New Zealand insect fossils are rare but a single fossil of a sooty mould cyclaxyrid was recently described from Cretaceous Burmese amber, a family that was previously known from two extant New Zealand species. Well-preserved fossils like this one are recasting Earth history, and, based on a wealth of additional specimens, we re-evaluate the taxonomy of Cretaceous cyclaxyrids and one Eocene species here transferred to Cyclaxyridae. Cyclaxyridae are highly tied to the sooty mould community and have now been discovered to occur in disparate biogeographic realms in deep time. Our discovery indicates that the family, and perhaps the sooty mould community in general, was widespread in Pangaea from at least the Cretaceous and survived as a relict in New Zealand. Persistence of a sooty mould ecosystem in New Zealand and fungal specialization may not necessarily be an evolutionary ‘dead-end’ for cyclaxyrids and other insects.
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.
Convergence between the Indian and Asian plates has reshaped large parts of Asia, changing regional climate and biodiversity, yet geodynamic models fundamentally diverge on how convergence was accommodated since the India–Asia collision. Here we report palaeomagnetic data from the Burma Terrane, which is at the eastern edge of the collision zone and is famous for its Cretaceous amber biota, to better determine the evolution of the India–Asia collision. The Burma Terrane was part of a Trans-Tethyan island arc and stood at a near-equatorial southern latitude at ~95 Ma, suggesting island endemism for the Burmese amber biota. The Burma Terrane underwent significant clockwise rotation between ~80 and 50 Ma, causing its subduction margin to become hyper-oblique. Subsequently, it was translated northward on the Indian Plate by an exceptional distance of at least 2,000 km along a dextral strike-slip fault system in the east. Our reconstructions are only compatible with geodynamic models involving an initial collision of India with a near-equatorial Trans-Tethyan subduction system at ~60 Ma, followed by a later collision with the Asian margin.
Significance
Aquatic organisms are rarely found in amber, but when they occur they provide invaluable evidence for the better understanding of amber taphonomy and past ecosystems. We report an ammonite and several marine gastropods alongside a mixed assemblage of intertidal and terrestrial forest floor organisms in mid-Cretaceous Burmese amber. Our discovery indicates that the Burmese amber forest was living near a dynamic and shifting coastal environment. The ammonite also provides supporting evidence for the age of the amber, which is still debated, and represents a rare example of dating using fossils present inside the amber.
The origin and early evolutionary history of polyphagan beetles have been largely based on evidence from the derived and diverse ‘core Polyphaga’, whereas little is known about the species-poor basal polyphagan lineages, which include Scirtoidea (Clambidae, Decliniidae, Eucinetidae, and Scirtidae) and Derodontidae. Here, we report two new species Acalyptomerus thayerae sp. nov. and Sphaerothorax uenoi sp. nov., both belonging to extant genera of Clambidae, from mid-Cretaceous Burmese amber. Acalyptomerus thayerae has a close affinity to A. herbertfranzi, a species currently occurring in Mesoamerica and northern South America. Sphaerothorax uenoi is closely related to extant species of Sphaerothorax, which are usually collected in forests of Nothofagus of Australia, Chile, and New Zealand. The discovery of two Cretaceous species from northern Myanmar indicates that both genera had lengthy evolutionary histories, originated at least by the earliest Cenomanian, and were probably more widespread than at present. Remarkable morphological similarities between fossil and living species suggest that both genera changed little over long periods of geological time. The long-term persistence of similar mesic microhabitats such as leaf litter may account for the 99 Myr morphological stasis in Acalyptomerus and Sphaerothorax. Additionally, the extinct staphylinoid family Ptismidae is proposed as a new synonym of Clambidae, and its only included species Ptisma zasukhae is placed as incertae sedis within Clambidae.
Burmese amber represents the world’s most diverse biota in the Mesozoic. Previous studies have focused on the biodiversity of its inclusions, as well as pholadid borings. Here we report a variety of marine animals symbiotic with or adhere to Burmese amber or the amber deposits, including crinoid columns, corals and oysters. We propose that there is no distinct evidence indicating the secondary transportation of Burmese amber over long distances. The ancient sedimentary environment was likely located in the coastal area. The hardening time of the resin was not long after secretion. The resin has been mixed with fragments of marine organisms in the ancient sediments, and has been deposited for a long time. The zircon age in the sediments surrounding amber approximately represents the age of Burmese amber, but due to limits of the method, the current zircon U-Pb SIMS age may be younger. Therefore, as far as the situation is concerned, the age of Burmese amber may be close to the boundary between the Albian and Cenomanian, or even late Albian. We suggest that it is plausible to generally refer to the age of Burmese amber as mid-Cretaceous, and a precise age requires further biostratigraphic and chronological studies.
Burmese amber is an extremely important source of mid-Cretaceous plant and animal remains with over 870 species of organisms, ranging from protozoa to vertebrates, described from this source. The amber mines are located on the West Burma Block that according to geologists was originally part of Gondwana. The present study introduces some angiosperms and insects in Burmese amber whose closest extant relatives have a Gondwanan distribution and there is no previous evidence of a Laurasian distribution. Based on this evidence, it is proposed that organisms in Burmese amber represent a selection of tropical to subtropical life forms that inhabited the interconnected continents of Gondwana in the Early Cretaceous. Based on the fossil record of angiosperms and their diversity in Burmese amber, the West Burma Block could not have rafted from Gondwana to SE Asia before the Early Cretaceous.
Through the lens of the fossil record, angiosperm diversification precipitated a Cretaceous Terrestrial Revolution (KTR) in which pollinators, herbivores and predators underwent explosive co‐diversification. Molecular dating studies imply that early angiosperm evolution is not documented in the fossil record. This mismatch remains controversial.
We used a Bayesian molecular dating method to analyse a dataset of 83 genes from 644 taxa and 52 fossil calibrations to explore the effect of different interpretations of the fossil record, molecular clock models, data partitioning, among other factors, on angiosperm divergence time estimation.
Controlling for different sources of uncertainty indicates that the timescale of angiosperm diversification is much less certain than previous molecular dating studies have suggested. Discord between molecular clock and purely fossil‐based interpretations of angiosperm diversification may be a consequence of false precision on both sides.
We reject a post‐Jurassic origin of angiosperms, supporting the notion of a cryptic early history of angiosperms, but this history may be as much as 121 Myr, or as little as 23 Myr. These conclusions remain compatible with palaeobotanical evidence and a more general KTR in which major groups of angiosperms diverged later within the Cretaceous, alongside the diversification of pollinators, herbivores and their predators.
Beetles (Coleoptera) are the most diverse and species-rich group of insects, and a robust, time-calibrated phylogeny is fundamental to understanding macroevolutionary processes that underlie their diversity. Here we infer the phylogeny and divergence times of all major lineages of Coleoptera by analyzing 95 protein-coding genes in 373 beetle species, including ~67% of the currently recognized families. The subordinal relationships are strongly supported as Polyphaga (Adephaga (Archostemata, Myxophaga)). The series and superfamilies of Polyphaga are mostly monophyletic. The species-poor Nosodendridae is robustly recovered in a novel position sister to Staphyliniformia, Bostrichiformia, and Cucujiformia. Our divergence time analyses suggest that the crown group of extant beetles occurred ~297 million years ago (Mya) and that ~64% of families originated in the Cretaceous. Most of the herbivorous families experienced a significant increase in diversification rate during the Cretaceous, thus suggesting that the rise of angiosperms in the Cretaceous may have been an 'evolutionary impetus' driving the hyperdiversity of herbivorous beetles.
The origin of the Cretaceous laminites of the Crato Formation, Araripe Basin, northeastern Brazil, has been intensely debated since the beginning of the last cen-tury. The monotonous, up to 10-m-thick succession com-posed of very fine fossiliferous laminites in the middle of the unit lacked diagnostic features for assigning a chemi-cally or biologically induced origin for this facies. The presence of a highly diverse and very well preserved alloch-thonous to parautochthonous fossil assemblage, associated with scattered halite pseudomorphs throughout the succes-sion, led many authors to believe that these limestones were chemically deposited in a highly stressful, evaporitic shal-low-water environment, such as a hypersaline lake close to marine environments. Recently, a micro- and ultrastruc-tural analysis of the laminites yielded structures undoubt-edly associated with a biological origin. Several examples of lithified in situ preserved coccoid and filamentous cells and extracellular polymeric substances suggest that the deposition of the laminated limestones was, at some levels, trongly influenced by microbial activity. Here, we record various examples of stromatolite microbialites (mounds, domes, and pseudo-columns) found at distinct stratigraphic levels in the middle part of the Crato Formation. Macro-, meso-, and microscopic features confirm the biologically induced mineralization and the existence of metabolic activity of microbes during the formation of the laminites. Biomat growth may also have played a major role in the excellent preservation of fossils in this famous Cretaceous Konservat-Lagerstätte from Brazil.
Resumo
A description of a new species of Cneoglossa (C. edsoni sp. n., type locality: Brazil, São Paulo State, City of São Paulo, Parque Estadual da Cantareira) is presented based on male and female adults, pupae and larvae. The synonymy of Buckodrillus Wittmer, 1948, with Cneoglossa Guérin-Méneville, 1843, is confirmed and C. brasiliensis (Wittmer, 1948), is a new combination. Larvae of C. edsoni were collected in submerged rotting brushwood in shallow flowing streams; pupae and adults were obtained from larvae reared in laboratory. Immatures of this genus were unknown up to date. Larval and/or adult features of species belonging to Anchytarsus Guérin-Méneville, 1843; Araeopidius Cockerell, 1906; Cladotoma Westwood, 1837; Epilichas White, 1859; Paralichas White, 1859; and, Ptilodactyla, Illiger, 1807 are described and illustrated. A cladistic analysis, conducted for 32 taxa of selected Byrrhoidea (sensu Lawrence & Newton, 1995) and 72 features of adults, larvae and pupae, is also given, in an attempt to clear up the systematic position of the Cneoglossidae. As a result of the analysis, Callirhipidae and Eulichadidae are excluded from Byrrhoidea (sensu Lawrence & Newton, 1995) and placed incertae sedis within the Elateriformia. The strict consensus cladogram of the 24 most parsimonious trees presented the following relationships: ((Byrrhidae ((Cneoglossidae, Psephenidae) ((Ptilodactylinae (Anchytarsinae, Epilichinae)) (Aploglossinae, Araeopidiinae (Cladotominae, Chelonariidae))))) (Larainae (Elminae (Lutrochidae (Hyphalinae (Limnichinae, Cephalobyrrhinae, Thaumastodinae, Dryopidae, Heteroceridae )))))). The monophyly of this assemblage is supported by three striking synapomorphies, the anterior process of metendosternite shortened, the proximal end of radial cell with acute inner angle, and the wing folding of the dryopoid-type. A sister-group relationship between Cneoglossidae and Psephenidae is indicated by the presence of paired dorsal glandular openings on abdominal tergites of adult, + five homoplasic features. Elmidae, Limnichidae and Ptylodactylidae, as currently defined, are paraphyletic. Ptylodactylidae is monophyletic with the inclusion of Chelonariidae.
Lasiosynidae n. fam. is proposed for the genera Lasiosyne Tan, Ren & Chih 2007 (transferred from Archostemata to Polyphaga), Anacapitis Yan 2009 Tarsomegamerus Zhang 2005 (proposed in the superfamily Chrysomeloidea) and Bupredactyla n. gen. The new family is regarded in composition of the infraorder Elateriformia without a more detailed attribution, because it demonstrates a mixture of characters of different families and superfamilies, i.e. somehow an intermediate position between the superfamilies Dascilloidea, Elateroidea, Buprestoidea and Byrrhoidea with most resemblance to Dascillidae, Schizopodidae, Eulichadidae, Ptilodactylidae and Callirrhipidae and probable more close relationship to the last three families. Four new fossil species of the genus Lasiosyne: L. daohugouensis n. sp., L. fedorenkoi n. sp., L gratiosa n. sp., L. quadricollis n. sp., and also Bupredactyla magna n. sp. are described from the Middle Jurassic Jiulongshan Formation of eastern Inner Mongolia, China. A probable generic composition of the new family is considered. The synonymy of generic names Anacapitis Yan 2009 and Brachysyne Tan & Ren 2009, n. syn. as well as synonymy of species names Lasiosyne euglyphea Tan, Ren & Chih 2007, Pappisyne eucallus Tan & Ren 2009, n. syn. and Pappisyne spathulata Tan & Ren 2009, n. syn. are proposed.
This is a supplement to the Burmese (Myanmar) amber checklist and bibliography covering taxa described or recorded during 2020, plus a few earlier records that were missed previously. Up to the end of 2020, 1,859 species were recorded from Kachin amber of which 362 were named in 2020, which is the highest number of species named from any kind of amber in one year. Two species were also named from older Hkamti amber.
Araripenepa vetussiphonis gen. et sp. nov., the oldest representative of the family Nepidae is described from the Early Cretaceous Crato Formation in Brazil. It is considered as the sister group of all the extant Nepidae, having retained tarsi with three tarsomeres and a transverse pronotum. The general habitus, elongate siphon, and strong grasping legs strongly suggest that its biology was very similar to that of the extant Nepa spp., viz. a predaceous bug living in the mud and/or the abundant aquatic vegetation of the Crato palaeolake. Possibly, the Triassic—Jurassic representatives of the stem group of Nepidae had no elongate siphon and were similar to Cratonepa, an enigmatic Nepoidea previously described from the Crato Formation. The development of the siphon was possibly favoured by the diversification of the lacustrine plants that occurred during the Early Cretaceous.
The Lower Cretaceous Crato Formation is a well-known Konservat Lagerstätte with a very rich entomofauna. The Odonata are especially very diverse and extensively studied (Bechly, 1998, 2000, 2007, 2010; Nel et al., 1998; Bechly et al., 2001; Bechly & Ueda, 2002) with representatives of all the extant anisopteran main subgroups. This fauna is especially interesting because it comprises some of the oldest and most ‘basal’ groups of the highly diverse extant ‘libelluloid’ dragonflies, or Clavilabiata Bechly, 1996. Among these, the monospecific family Araripephlebiidae Bechly, 1998 is remarkable in the highly specialized hind wing cubito-anal area that contains a curious supplementary longitudinal vein more or less parallel to AA and CuA, unique among the Odonata. Nevertheless, this family remained rather poorly known by three described specimens, only females.
Amber in Lebanon is found in more than 450 outcrops. It constitutes the oldest amber with intensive biological inclusions and is considered among the most important material enabling the knowledge of continental palaeobiodiversity from the very important Lower Cretaceous, a crucial period for the coevolution between flowering plants (angiosperms) and insects. This period is largely admitted to witnessing the first occurrence and early evolution of angiosperms. Most times biological inclusions in Lebanese amber represent records of the earliest representatives of modern living insect families or the youngest ones for extinct families. Latest literature, geological data on age and lists of amber outcrops (yielding fossil inclusions), and described taxa from Lebanese amber are given.
This is a supplement to Ross (2019) covering all taxa described or recorded in Burmese amber during 2019, plus a few earlier records that were missed previously. Up to the end of 2019, 1,478 species were recorded from Burmese (Kachin) amber of which 276 were named or recorded in 2019.
Variegated mud-loving beetles, or Heteroceridae, are a small family belonging to the polyphagan superfamily Byrrhoidea. To date, only two poorly preserved compression fossils have been known from the Early Cretaceous of Eurasia. Here we describe the first heterocerid beetles in mid-Cretaceous amber from northern Myanmar, Excavotarsus lini gen. et sp. nov. and Ex. minor sp. nov.. The two new fossil species are distinguished from all extant heterocerids by their elongate body shape, apical 8–9 antennomeres forming a loose serrate club, pronotum longer than wide, protibia lacking robust spines, and two-segmented protarsi. This peculiar combination of plesiomorphic and derived characters suggests that Excavotarsus represents an early-diverging lineage of Heteroceridae and indicates that the family originated and diversified before the mid-Cretaceous.
Three fossil species classified in the family Monotomidae are described from mid-Cretaceous Burmese amber. They are attributed to the recent tribe Lenacini Crowson, known so far from a single extant endemic species, Lenax mirandus Sharp, from New Zealand. Lenax karenae Liu, Tihelka, McElrath and Yamamoto sp. nov., Cretolenax carinatus Liu, Tihelka, McElrath and Yamamoto gen. et sp. nov., and Cretolenax diabolus Tihelka, Liu, McElrath and Yamamoto gen. et sp. nov. reveal high diversity and a much broader distribution of Lenacini during the Cretaceous. The discovery of the Burmese amber taxa closely related to the New Zealand endemic supports growing evidence about the Gondwanan origin of the West Burma Block.
Adults of Geoparnus rhinoceros sp. nov. (Dryopidae) are described from Borneo (Sarawak, Malaysia). The male of the new species possesses a distinct horn-like process on the clypeus, a character, which has so far not been reported from Dryopidae. The type material was collected in primary rain forest by sifting forest floor debris. Analysis of variance of metric characters was performed.
A list of all known taxa described or recorded from Burmese amber from the published literature up to the end of 2018 is given, along with a comprehensive bibliography. The history of the study of inclusions is summarised, and demonstrates that the number of species has risen exponentially over the past two decades. The first three species were named in 1916 and by the end of 1920 a total of 42 species had been named by T.D.A. Cockerell. Only three more species were named by 1999 though by the end of 2018 the total had risen to an incredible 1,192 species, of which over half were named in the past three years. Some 320 species were named in 2018, the highest number described from one type of amber in any one year in the entire history of amber studies.
A molecular phylogeny of Ptilodactylidae shows that Podabrocephalus Pic, 1913, the type genus of Podabrocephalidae Pic, 1930 is closely related to Ptilodactyla Illiger, 1807 and other genera of Ptilodactylinae. Consequently, Podabrocephalidae Pic, 1930 syn. n. is proposed as a junior synonym of Ptilodactylidae Laporte, 1836. Earlier authors used the highly modified morphology of the male to justify a high rank for Podabrocephalus. The molecular phylogeny of Ptilodactylidae further indicates that Paralichas White, 1859 (Cladotominae) does not form a monophylum with remaining ptilodactylids. Ptilodactylinae and an undescribed lineage from Indonesia are sister to the broadly delimited Anchytarsinae. Within Ptilodactylinae, Pherocladus Fairmaire, 1881 is sister to a clade formed by Ptilodactyla spp. and Podabrocephalus. We remove Falsotherius Pic, 1913 from Ptilodactylinae to Ptilodactylidae incertae sedis, and return Daemon Laporte, 1836 from Ptilodactylinae to Anchytarsinae. Cross validation of morphology- and DNA-based phylogenies is needed for interpreting phylogenetic inference in morphologically modified lineages.
Clavate (club-shaped) structures rimming mid-Cretaceous Burmese amber from Myanmar, previously misdiagnosed as fungal sporocarps, are shown to be domichnia (crypts) of martesiine bivalves (Pholadidae: Martesiinae). They are similar in form to Teredolites clavatus Leymerie, 1842 and Gastrochaenolites lapidicus Kelly & Bromley, 1984; however, the former identification is preferable, given that they are martesiine crypts in amber as opposed to a lithic substrate. Cross-cutting relationships between the clavate features and inclusions in the amber demonstrate that the features post-date hardening of the resin. The fills of the crypts are variable, including sand grade sediment of very fine to coarse sand grainsize, and sparry calcite cements. In some cases, the articulated valves of the pholadid bivalve responsible are visible inside the borings. However, one remarkable specimen contains two pairs of articulated shells ‘floating’ in amber, not associated with crypts; an observation that suggests that the resin was still liquid or soft when the bivalves were trapped in the resin. One individual is associated with an irregular sediment-filled feature and shows shell breakage. Formation of a solid rim around a liquid central volume has been documented in subaqueous bodies of resin in modern swamp forests, and argues for a close proximity between the amber-producing trees and a brackish water habitat for the bivalves. The presence of pyrite as thin films and crystal groups within Burmese amber is further consistent with such a depositional environment. Comparison of the size of pholadid body fossils with growth rates of modern equivalents allows the duration of boring activities to be estimated and suggests that small fossil pholadids in Burmese amber became trapped and died within 1–2 weeks of having settled on the resin. Larger examples present within well-formed domichnia formed in hardened resin. Since ‘hardground’ describes early lithified sediment as a substrate and ‘woodground’ describes wood as a substrate, the term ‘amberground’ is used here to described borings in an amber substrate.
One of the lasting controversies in phylogenetic inference is the degree to which specific evolutionary models should influence the choice of methods. Model-based approaches to phylogenetic inference (likelihood, Bayesian) are defended on the premise that without explicit statistical models there is no science, and parsimony is defended on the grounds that it provides the best rationalization of the data, while refraining from assigning specific probabilities to trees or character-state reconstructions. Authors who favour model-based approaches often focus on the statistical properties of the methods and models themselves, but this is of only limited use in deciding the best method for phylogenetic inference—such decision also requires considering the conditions of evolution that prevail in nature. Another approach is to compare the performance of parsimony and model-based methods in simulations, which traditionally have been used to defend the use of models of evolution for DNA sequences. Some recent papers, however, have promoted the use of model-based approaches to phylogenetic inference for discrete morphological data as well. These papers simulated data under models already known to be unfavourable to parsimony, and modelled morphological evolution as if it evolved just like DNA, with probabilities of change for all characters changing in concert along tree branches. The present paper discusses these issues, showing that under reasonable and less restrictive models of evolution for discrete characters, equally weighted parsimony performs as well or better than model-based methods, and that parsimony under implied weights clearly outperforms all other methods.
Although both the Yanliao and Jehol vertebrate assemblages are known for exceptional preservation of feathered dinosaurs, mammals, pterosaurs, lizards, salamanders, and fish, the Early Cretaceous Jehol Biota also contains birds, choristoderes, frogs, and turtles that are currently lacking in the Jurassic Yanliao Biota. The vertebrate assemblages of the Yanliao and Jehol biotas are very distinct from each other, with the salamander Liaoxitriton being the sole unquestionably shared taxon on generic level. Both assemblages contain mainly stem relatives of major clades of extant vertebrates, with all genera and species extinct. Crown group taxa are restricted to some fishes, salamanders, frogs, and turtles.
Coleoptera: Elmidae and Protelmidae includes 151 extant genera and 1501 extant species as well as four fossil species described before 2015. Protelmidae are here elevated from tribal rank to family rank.
The superfamilies of Elateriformia have been in a state of flux since their establishment. The recent classifications recognize Dascilloidea, Buprestoidea, Byrrhoidea and Elateroidea. The most problematic part of the elateriform phylogeny is the monophyly of Byrrhoidea and the relationships of its families. To investigate these issues, we merged more than 500 newly produced sequences of 18S rRNA, 28S rRNA, rrnL mtDNA and cox1 mtDNA for 140 elateriform taxa with data from GenBank. We assembled an all-taxa (488 terminals) and a pruned data set, which included taxa with full fragment representation (251 terminals); both were aligned in various programs and analysed using maximum-likelihood criterion and Bayesian inference. Most analyses recovered monophyletic superfamilies and broadly similar relationships; however, we obtained limited statistical support for the backbone of trees. Dascilloidea were sister to the remaining Elateriformia, and Elateroidea were sister to the clade of byrrhoid lineages including Buprestoidea. This clade mostly consisted of four major lineages, that is (i) Byrrhidae, (ii) Dryopidae + Lutrochidae, (iii) Buprestoidea (Schizopodidae sister to Buprestidae) and (iv) a clade formed by the remaining byrrhoid families. Buprestoidea and byrrhoid lineages, with the exception of Byrrhidae and Dryopidae + Lutrochidae, were usually merged into a single clade. Most byrrhoid families were recovered as monophyletic. Callirhipidae and Eulichadidae formed independent terminal lineages within the Byrrhoidea-Buprestoidea clade. Paraphyletic Limnichidae were found in a clade with Heteroceridae and often also with Chelonariidae. Psephenidae, represented by Eubriinae and Eubrianacinae, never formed a monophylum. Ptilodactylidae were monophyletic only when Paralichas (Cladotominae) was excluded. Elmidae regularly formed a clade with a bulk of Ptilodactylidae; however, elmid subfamilies (Elminae and Larainae) were not recovered. Despite the densest sampling of Byrrhoidea diversity up to date, the results are not statistically supported and resolved only a limited number of relationships. Furthermore, questions arose which should be considered in the future studies on byrrhoid phylogeny.
Version 1.5 of the computer program TNT completely integrates landmark data into phylogenetic analysis. Landmark data consist of coordinates (in two or three dimensions) for the terminal taxa; TNT reconstructs shapes for the internal nodes such that the difference between ancestor and descendant shapes for all tree branches sums up to a minimum; this sum is used as tree score. Landmark data can be analysed alone or in combination with standard characters; all the applicable commands and options in TNT can be used transparently after reading a landmark data set. The program continues implementing all the types of analyses in former versions, including discrete and continuous characters (which can now be read at any scale, and automatically rescaled by TNT). Using algorithms described in this paper, searches for landmark data can be made tens to hundreds of times faster than it was possible before (from T to 3T times faster, where T is the number of taxa), thus making phylogenetic analysis of landmarks feasible even on standard personal computers.
The “Grès du Liban” [Sandstone of Lebanon] is the basal lithostratigraphic unit for the Cretaceous series in Lebanon. In the upper part of these siliciclastic-dominated strata we identified three discrete intervals characterized by their richness in amber with biological inclusions, mostly insects. The middle and upper intervals previously attributed to an Early Aptian (= Bedoulian) age are nowadays ascribed to the Early and Late Barremian respectively; the lower interval is Early Barremian or possibly older. Besides that it is suggested that pieces of amber with inclusions from the middle and upper intervals could be reworked from the lower interval. In conclusion, the new dating of arthropod-bearing localities allows us to push back in time (at least to the Early Barremian) the first occurrences of all biological inclusions found therein. © 2016 Elsevier B.V. and Nanjing Institute of Geology and Palaeontology, CAS
This beautifully illustrated 2007 volume describes the entire flora and fauna of the famous Lower Cretaceous Crato Formation of Brazil - one of the world's most important fossil deposits, exhibiting exceptional preservation. A wide range of invertebrates and vertebrates are covered, including extended sections on pterosaurs and insects. Two chapters are devoted to plants. Many of the chapters include descriptions of new species and re-descriptions and appraisals of taxa published in obscure places, rendering them available to a wider audience. Fossil descriptions are supported by detailed explanations of the geological history of the deposit and its tectonic setting. Drawing on expertise from around the world and specimens from the most important museum collections, this book forms an essential reference for researchers and enthusiasts with an interest in Mesozoic fossils.
This paper overviews more than 39 families of fossil Coleoptera from Lower Cretaceous Lebanese amber from nine outcrops. Lebanese amber contains the oldest representatives of the families Scydmaenidae (considered by some as a subfamily of Staphylinidae), Ptiliidae, Elodophalmidae, Clambidae, Throscidae, Lebanophytidae fam. n., Ptilodactylidae, Cantharidae, Melyridae, Dasytidae, Dermestidae, Ptinidae, Kateretidae, Erotylidae, Latridiidae, Laemophloeidae, Salpingidae, Anthicidae, Melandryidae, Aderidae, Curculionidae (Scolytinae). The families Chelonariidae and Scraptiidae are known from both Lebanese amber and Baissa, with both sites having a comparable age. The subfamilies Trechinae (Carabidae), Euaesthetinae (Staphylinidae) and Liparochrinae (Hybosoridae) first appear in the fossil record in Lebanese amber. The Coleoptera in Lebanese amber mostly belong to groups with arboreal habits (as found today in wood and tree fungi). Eochelonarium belle gen. et sp. n., Rhizophtoma synchrotronica sp. n., Rhizobactron marinae gen et sp. n. and Atetrameropsis subglobosa gen. et sp. n. are described from Lebanese amber. A new subfamily in the family Cerophytidae is proposed for Aphytocerus communis Zherichin, 1977 (Aphytocerinae subfam. n.) and new genus Baissopsis gen.nov. is erected for Baissophytum amplus Chang, Kirejtshuk et Ren, 2011. Also a new interpretation of the taxon “Lasiosynidae” is provided by placing it as a subfamily in the family Eulichadidae with two genera (Lasiosyne Tan, Ren et Shih, 2007 and Bupredactyla Kirejtshuk, Chang, Ren et Shih, 2010), while the other genera initially regarded as “Lasiosynidae” were tentatively transferred into Eulichadinae sensu n. (Mesodascilla Martynov, 1926; Tarsomegamerus Zhang, 2005; Brachysyne Tan et Ren, 2009; Anacapitis Yan, 2009; Parelateriformius Yan et Wang, 2010 and Cretasyne Yan, Wang et Zhang, 2013) with the new synonymy of Tarsomegamerus and Parelateriformius syn. n. The genus Mesaplus Hong, 1983 described in the family Triaplidae is also transvered to Eulichadinae. The genera Artematopodites Ponomarenko, 1990; Dzeregia Ponomarenko, 1985 and Glaphyropteroides Handlirsch, 1906 proposed for species known only by separate elytra and recently included in the “family” Lasiosynidae (Yan et al., 2013) are regarded as Elateriformia incertae sedis. The first insect from the newly discovered outcrops of Nabaa Es-Sukkar – Brissa: Caza (District) Sir Ed-Danniyeh, Mouhafazet (Governorate) Loubnan Esh-Shimali (North Lebanon) is described and the first general description of this outcrop is made.
Significance
We report on the unique discovery of Jurassic and Cretaceous carrion beetles (Silphidae) from China and Myanmar, early relatives of one of the most protected of beetle species in North America, and which clearly preserve evidence indicative of complex parental care. This finding represents the earliest evidence of parental care, a behavioral repertoire that is the first step in the development of truly social behavior and one that is intensely studied by ecologists, ethologists, and evolutionary biologists alike. Our fossils clearly span the origins of parent–offspring communication and allow us to provide a robust estimate of the time of origin for this complex behavior.