Michael S. Engel

"This book chronicles, for the first time, the complete evolutionary history of insects: their living diversity, relationships, and 400 million years of fossils. Whereas other volumes have focused on either living species or fossils, this is the first comprehensive synthesis of all aspects of insect evolution. Current estimates of phylogeny are used to interpret the 400-million-year fossil record of insects, their extinctions, and radiations." "Evolution of the Insects is beautifully illustrated with more than 900 photo- and electron micrographs, drawings, diagrams, and field photographs, many in full color and virtually all original. The book will appeal to anyone engaged with insect diversity: professional entomologists and students, insect and fossil collectors, and naturalists."--BOOK JACKET.

Amber from Kachin, northern Burma, has been used in China for at least a millennium for carving decorative objects, but the only scientific collection of inclusion fossils, at the Natural History Museum, London (NHML), was made approximately 90 years ago. Age of the material was ambiguous, but probably Cretaceous. Numerous new records and taxa occur in this amber, based on newly excavated material in the American Museum of Natural History (AMNH) containing 3100 organisms. Without having all groups studied, significant new records and taxa thus far include the following (a † refers to extinct taxa): For Plants: An angiosperm flower (only the third in Cretaceous amber), spores and apparent sporangia of an unusual but common fungus, hepatophyte thalli and an archegoniophore of Marchantiaceae, and leafy shoots of Metasequoia (Coniferae). Metasequoia is possibly the source of the amber. For Animals: Mermithidae and other Nematoda; the oldest ixodid tick (a larval Amblyomma); bird feathers; and the only Mesozoic record of the Onychophora (“velvet” worms), described as †Cretoperipatus burmiticus, n. gen., n. sp. (Peripatidae). Poinar's classification of the Onychophora is substantially revised. Still largely unstudied, the fauna of mites (Acari) and spiders (Araneae) appears to be the most diverse ones known for the Mesozoic. For Insecta: Odonata indet. (wing fragment); Plecoptera indet.; new genera of Dermaptera, Embiidina, and Zoraptera (the latter two as the only definitive Mesozoic fossils of their orders). Within Hemiptera, there are primitive new genera in the Aradidae, Hydrometridae, Piesmatidae, Schizopteridae, and Cimicomorpha (Heteroptera), as well as in †Tajmyraphididae (Aphidoidea), and †′otopsyllidiidae. An adult snakefly (Raphidioptera: †Mesoraphidiidae) is the smallest species in the order, and new genera occur in the Neuroptera: Coniopterygidae, Berothidae, and Psychopsidae, as well as larvae of apparent Nevrorthidae. Coleoptera are largely unstudied, but are probably the most diverse assemblage known from the Cretaceous, particularly for Staphylinidae. An adult lymexylid, the most primitive species of Atractocerus, is the first Mesozoic record of the family. In Hymenoptera there are primitive ants (Formicidae: Ponerinae n. gen., and †Sphecomyrma n.sp [Sphecomyrminae]), the oldest record of the Pompilidae, and significant new records of †Serphitidae and †Stigmaphronidae, among others. Diptera are the most diverse and abundant, with the oldest definitive Blephariceridae and mosquito (Culicidae), as well as new genera in the Acroceridae, Bibionidae, Empidoidea; a new genus near the enigmatic genus Valeseguya, and an unusual new genus in the †Archizelmiridae. †Chimeromyia (Diptera: Eremoneura), known previously in ambers from the Lower Cretaceous, is also represented.The stratigraphic distribution of exclusively Mesozoic arthropods in Burmese amber is reviewed, which indicates a probable Turonian-Cenomanian age of this material (90–100 Ma). Paleofaunal differences between the NHML and AMNH collections are discussed, as is the distinct tropical nature of the original biota. Burmese amber probably harbors the most diverse biota in amber from the Cretaceous, and one of the most diverse Mesozoic microbiotas now known.

Abstract The bees found as Baltic amber inclusions are revised and the history of studies on these fossils is briefly reviewed. In total this subtropical Eocene fauna contains 36 species and 18 genera, all extinct. These are classified here into nine tribes, six subfamilies, and five families of which six tribes and one family are unknown in the modern fauna. The following taxa are described as new to science: PALEOMELITTIDAE, new family; Boreallodapini, Electrobombini, Eomacropidini, Melikertini, Protolithurgini, new tribes; Ctenoplectrellina, new subtribe; Boreallodape, Electrobombus, Electrolictus, Eomacropis, Glaesosmia, Liotrigonopsis, Melissites, Paleomelitta, Protolithurgus, Succinapis, Thaumastobombus, new genera; Boreallodape baltica, B. mollyae, B. striebichi, Ctenoplectrella cockerelli, C. grimaldii, Electrapis krishnorum, Electrobombus samlandensis, Electrolictus antiquus, Eomacropis glaesaria, Glaesosmia genalis, Glyptapis densopunctata, G. disareolata, Liotrigonopsis rozeni, Melikertes clypeatus, Melissites trigona, Paleomelitta nigripennis, Protobombus basilaris, Protolithurgus ditomeus, Succinapis goeleti, S. micheneri, S. proboscidea, Thaumastobombus andreniformis, new species (seven new family-, 11 new genus-, and 22 new species-group taxa). The genus Electrapis is found to be paraphyletic and the subgenera Melikertes, Roussyana, and Protobombus are given generic status outside of Electrapis. The subtribe Electrapina is elevated to tribal rank among the corbiculate Apinae and the subfamily Glyptapinae of Cockerell is reduced to subtribal rank within Osmiini. The genera Chalcobombus and Sophrobombus are newly synonymized with Protobombus. Glyptapis reducta Cockerell is synonymized with G. fuscula Cockerell, Electrapis minuta Kelner-Pillault with Apis palmnickenensis Roussy, Ctenoplectrella splendens Kelner-Pillault and C. dentata Salt both with C. viridiceps Cockerell, Electrapis apoides Manning and Chalcobombus humilis Cockerell both with Protobombus indecisus Cockerell, and Electrapis bombusoides Kelner-Pillault with E. tornquisti Cockerell (new synonymies). The following new combinations are proposed: Electrapis martialis (Cockerell), Melikertes proavus (Menge), M. stilbonotus (Engel), Kelneriapis eocenica (Kelner-Pillault), Protobombus fatalis (Cockerell), P. hirsutus (Cockerell), and Electrapis martialis (Cockerell) (new combinations). A lectotype is designated for Electrapis minuta Kelner-Pillault and neotypes designated for Apis meliponoides Buttel-Reepen, A. palmnickenensis Roussy, Chalcobombus humilis Cockerell, C. hirsutus Cockerell, C. martialis Cockerell, Ctenoplectrella dentata Salt, C. viridiceps Cockerell, Electrapis tornquisti Cockerell, Glyptapis reticulata Cockerell, G. neglecta Salt, Protobombus indecisus Cockerell, P. tristellus Cockerell, and Sophrobombus fatalis Cockerell. The subfamily Xylocopinae is recorded for the first time from amber, and the families Halictidae and Melittidae are confirmed as occurring in Baltic amber. The oldest fossils of the Halictidae, Megachilidae (Lithurginae and Megachilinae), Melittidae, and Xylocopinae are reported and described herein. Keys are presented for the identification of the Baltic amber bees. Three new recent taxa are also proposed: Penapini, Redivivini, and Meliturgulini (new tribes). Cladistic analyses of the Lithurginae, Xylocopinae, and corbiculate Apinae are presented. Preliminary investigation of lithurgine relationships demonstrates that the fossil, Protolithurgus, is sister to extant genera of the subfamily and that Lithurgus s.s. is likely paraphyletic with respect to Lithurgopsis and Microthurge. Xylocopine relationships are generally in accord with those previously proposed: Xylocopini sister to all other tribes and Manueliini sister to the abruptly narrowed mandible tribes (i.e., Ceratinini, Allodapini, and Boreallodapini). Boreallodapini, despite some apparently apomorphic similarities with Ceratinini, is supported as sister to Allodapini. The paleontological evidence for corbiculate bee phylogeny reinforces traditional concepts over relationships of these tribes and contradicts recent molecular studies. Two extinct corbiculate tribes are based on specimens that exhibit worker morphologies indicative of advanced eusocial behavior and are related to the living, advanced eusocial tribes Apini and Meliponini, supporting a hypothesis of a single origin for this behavioral characteristic. The composition of the Eocene bee fauna of Europe is discussed. The fauna is predominantly composed of long-tongued bees, but some short-tongued bees are represented in the families Halictidae, Paleomelittidae, and Melittidae. Bees in Baltic amber are generally allied with groups currently inhabiting sub-Saharan Africa and southeast Asia, a biogeographic pattern common to many Baltic amber groups. The phylogeny and origin of bees are discussed. The bees are a derived, monophyletic group of the spheciform wasps and presumably arose sometime in the earliest mid-Cretaceous after the origin of angiosperms. Reports of bees from Jurassic strata or earlier are all refuted. All available evidence supports the idea that bees originated shortly after flowering plants and diversified into higher lineages contemporaneously with the radiations of angiosperms. Major bee lineages (i.e., families) were thus presumably established by the Late Cretaceous. A catalog of bees presently known in amber and copal is appended as is a catalog of family- and genus-group names for fossil bees (both amber inclusions and compression fossils). A preliminary outline of the tribal classification of Recent and fossil bees with their geological distribution is presented (with three new family-group taxa for living bees). ZUSAMMENFASSUNG Die als Inklusen des Baltischen Bernsteins nachgewiesenen Bienen werden revidiert und ihre Erforschungsgeschichte kurz dargestellt. Insgesamt umfasst diese subtropische Fauna des Eozäns 36 Arten und 18 Gattungen, die alle ausgestorben sind. Diese werden in neun Tribus, sechs Unterfamilien, und fünf Familien klassifiziert, von denen sechs Tribus und eine Familie in der heutigen Fauna unbekannt sind. Die folgenden Taxa sind neu für die Wissenschaft: PALEOMELITTIDAE, fam. nov.; Boreallodapini, Electrobombini, Eomacropidini, Melikertini, Protolithurgini, tribus nov.; Ctenoplectrellina, subtribus nov.; Boreallodape, Electrobombus, Electrolictus, Eomacropis, Glaesosmia, Liotrigonopsis, Melissites, Paleomelitta, Protolithurgus, Succinapis, Thaumastobombus, gen. nov.; Boreallodape baltica, B. mollyae, B. striebichi, Ctenoplectrella cockerelli, C. grimaldii, Electrapis krishnorum, Electrobombus samlandensis, Electrolictus antiquus, Eomacropis glaesaria, Glaesosmia genalis, Glyptapis densopunctata, G. disareolata, Liotrigonopsis rozeni, Melikertes clypeatus, Melissites trigona, Paleomelitta nigripennis, Protobombus basilaris, Protolithurgus ditomeus, Succinapis goeleti, S. micheneri, S. proboscidea, Thaumastobombus andreniformis, spec. nov. (sieben neue Familien-, 11 neue Gattungs-, und 22 neue Artengruppentaxa). Die Gattung Electrapis ist paraphyletisch, und die Untergattungen Melikertes, Roussyana, und Protobombus werden unabhängig von Electrapis als Gattungen akzeptiert. Die Untertribus Electrapina erhält den Rang einer Tribus innerhalb der corbiculaten Apinae, und die Unterfamilie Glyptapinae von Cockerell wird als Untertribus innerhalb der Osmiini interpretiert. Die Gattungen Chalcobombus und Sophrobombus werden neu mit Protobombus synonymisiert. Folgende Arten werden miteinander synonymisiert: Glyptapis reducta Cockerell mit G. fuscula Cockerell, Electrapis minuta Kelner-Pillault mit Apis palmnickenensis Roussy, Ctenoplectrella splendens Kelner-Pillault und C. dentata Salt beide mit C. viridiceps Cockerell, Electrapis apoides Manning und Chalcobombus humilis Cockerell beide mit Protobombus indecisus Cockerell, Electrapis bombusoides Kelner-Pillault mit E. tornquisti Cockerell (neue Synonyme). Die folgenden Kombinationen werden neu eingeführt: Electrapis martialis (Cockerell), Melikertes proavus (Menge), M. stilbonotus (Engel), Kelneriapis eocenica (Kelner-Pillault), Protobombus fatalis (Cockerell), P. hirsutus (Cockerell), und Electrapis martialis (Cockerell) (neue Kombinationen). Ein Lectotypus wird für Electrapis minuta Kelner-Pillault festgelegt. Für folgende Arten werden Neotypen designiert: Apis meliponoides Buttel-Reepen, A. palmnickenensis Roussy, Chalcobombus humilis Cockerell, C. hirsutus Cockerell, C. martialis Cockerell, Ctenoplectrella dentata Salt, C. viridiceps Cockerell, Electrapis tornquisti Cockerell, Glyptapis reticulata Cockerell, G. neglecta Salt, Protobombus indecisus Cockerell, P. tristellus Cockerell, und Sophrobombus fatalis Cockerell. Die Unterfamilie Xylocopinae wird erstmals in Bernstein nachgewiesen, und das Vorkommen der Familien Halictidae und Melittidae in Baltischem Bernstein wird bestätigt. Die ältesten Fossilien der Halictidae, Megachilidae (Lithurginae und Megachilinae), Melittidae, und Xylocopinae werden nachgewiesen und beschrieben. Bestimmungsschlüssel für die Bienen des Baltischen Bernsteins werden erstellt. Drei rezenten Taxa sind neu: Penapini, Redivivini, und Meliturgulini (tribus nov.). Kladistische Analysen der Lithurginae, Xylocopinae, und der corbiculaten Apinae werden vorgestellt. Vorläufige Untersuchungen der verwandtschaftlichen Beziehungen der Lithurginae zeigen, dass die fossile Protolithurgus Schwestergruppe der rezenten Gattungen der Lithurginae ist. Außerdem ist Lithurgus s.s. wahrscheinlich paraphyletisch in Bezug auf Lithurgopsis und Microthurge. Die Verwandtschaftsbeziehungen der Xylocopinae stimmen grundsätzlich mit den bisher angenommenen überein: Die Xylocopini sind die Schwestergruppe aller übrigen Tribus, während die Manueliini die Schwestergruppe aller Tribus mit Arten mit abrupt verschmälerten Mandibeln ist (z.B. Ceratinini, Allodapini, und Boreallodapini). Boreallodapini wird a

Engel, Michael S, Ceríaco, Luis M P, Daniel, Gimo M, Dellapé, Pablo M, Löbl, Ivan, Marinov, Milen, Reis, Roberto E, Young, Mark T, Dubois, Alain, Agarwal, Ishan, Lehmann A., Pablo, Alvarado, Mabel, Alvarez, Nadir, Andreone, Franco, Araujo-Vieira, Katyuscia, Ascher, John S, Baêta, Délio, Baldo, Diego, Bandeira, Suzana A, Barden, Phillip, Barrasso, Diego A, Bendifallah, Leila, Bockmann, Flávio A, Böhme, Wolfgang, Borkent, Art, Brandão, Carlos R F, Busack, Stephen D, Bybee, Seth M, Channing, Alan, Chatzimanolis, Stylianos, Christenhusz, Maarten J M, Crisci, Jorge V, D'elía, Guillermo, Da Costa, Luis M, Davis, Steven R, De Lucena, Carlos Alberto S, Deuve, Thierry, Fernandes Elizalde, Sara, Faivovich, Julián, Farooq, Harith, Ferguson, Adam W, Gippoliti, Spartaco, Gonçalves, Francisco M P, Gonzalez, Victor H, Greenbaum, Eli, Hinojosa-Díaz, Ismael A, Ineich, Ivan, Jiang, Jianping, Kahono, Sih, Kury, Adriano B, Lucinda, Paulo H F, Lynch, John D, Malécot, Valéry, Marques, Mariana P, Marris, John W M, Mckellar, Ryan C, Mendes, Luis F, Nihei, Silvio S, Nishikawa, Kanto, Ohler, Annemarie, Orrico, Victor G D, Ota, Hidetoshi, Paiva, Jorge, Parrinha, Diogo, Pauwels, Olivier S G, Pereyra, Martín O, Pestana, Lueji B, Pinheiro, Paulo D P, Prendini, Lorenzo, Prokop, Jakub, Rasmussen, Claus, Rödel, Mark-Oliver, Rodrigues, Miguel Trefaut, Rodríguez, Sara M, Salatnaya, Hearty, Sampaio, Íris, Sánchez-García, Alba, Shebl, Mohamed A, Santos, Bruna S, Solórzano-Kraemer, Mónica M, Sousa, Ana C A, Stoev, Pavel, Teta, Pablo, Trape, Jean-François, Dos Santos, Carmen Van-Dúnem, Vasudevan, Karthikeyan, Vink, Cor J, Vogel, Gernot, Wagner, Philipp, Wappler, Torsten, Ware, Jessica L, Wedmann, Sonja, Zacharie, Chifundera Kusamba (2021): EDITORIAL The taxonomic impediment: a shortage of taxonomists, not the lack of technical approaches. Zoological Journal of the Linnean Society 193 (2): 381-387, DOI: 10.1093/zoolinnean/zlab072, URL: https://academic.oup.com/zoolinnean/article/193/2/381/6374389

Like ants, termites are entirely eusocial and have profound ecological significance in the tropics. Following upon recent studies reporting more than a quarter of all known fossil termites, we present the first phylogeny of termite lineages using exemplar Cretaceous, Tertiary, and Recent taxa. Relationships among Recent families were largely unaffected by the addition of extinct taxa, but the analysis revealed extensive grades of stem-group taxa and the divergence of some modern families in the Cretaceous. Rhinotermitidae, Serritermitidae, and the “higher” termites (family Termitidae), which comprise 84% of the world termite species, diverged and radiated entirely in the Tertiary, corresponding to a significant increase in termite individuals in the fossil record. Radiation of the higher termites may have affected the formation of terrestrial carbon reserves like oil and coal. The higher classification of Isoptera is slightly revised based on the phylogenetic results. The following new taxa are proposed: Cratomastotermitidae, new family; Euisoptera, new clade; Archotermopsidae, new family; and Neoisoptera, new clade. In addition, the families Stolotermitidae, Stylotermitidae, and Archeorhinotermitidae are newly recognized or resurrected, and the families Termopsidae and Hodotermitidae are significantly restricted in composition.