DOCSLIB.ORG

A Brief Guide to the Evolution of Flying Insects

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
A Brief Guide to the Evolution of Flying Insects Brief Guide Brief A Brief Guide to the evolution of flying insects Sabrina Simon s Sackler Institute for Comparative Genomics, American Museum of Natural History, New York, NY, USA Corresponding author: Sabrina Simon E-mail: [email protected] The evolutionary novelty – insect wings – ignited the mega-diversity and evolutionary success of insects and is one of the most fascinating issues in developmental and evolutionary biology. However, neither the origin of insect wings nor the early evolution of winged insects (Pterygota) is well understood. Origin of insect wings The origin of insect wings can be explained by two main hypotheses: insect wings represent a novel structure (Paranotal hypothesis) or a modification of an existing structure (Limb branch hypothesis) [1]. Based on the Paranotal hypothesis, wings evolved as novel outgrowths of the thoracic tergum, or paranotal lobe, as an adaption to gliding prior to the appearance of proper wings [2] (Figure 1a). The second hypothesis proposes that wings Figure 1. a) Colored: shown are the proposed structural homologies according to the two main hypotheses for the evolutionary origin of are a modification of limb branches that were insect wings. Based on the Paranotal hypothesis the paranotal lobes present in ancestral branched appendages of their (non-articulated) existing in the extinct Paleodictyoptera are homologous to the functional wings in modern insects. In contrast, ancestors and served as respiratory surfaces the Limb branch hypothesis considers the gills observed in mayfly (gills) before modification into powerful flight nymphs for respiration as homologous structures to insect wings. b) Proposed phylogenetic relationships for the earliest branching organs [3]. lineages of Pterygota: Pala, Palaeoptera; Meta, Metapterygota; and Both hypotheses are supported by developmental Chias, Chiastomyaria hypothesis. studies of regulatory genes but have been ques- tioned for various reasons. The Paranotal hypo- thesis is implausible in a parsimonious evolu- Keywords: insect evolution; systematics; wings; wing folding Please note: this contribution was not peer-reviewed. 1 tionary scenario – the derivation of mobile Neoptera (all other extant insects with wing flexion) structures (flapping wings) from primitively (Figure 1b). Here, a major problem is the immobile paranotal lobes. This suggests assignment of homology between the primarily secondarily derived wing articulation but the origin wingless outgroup (Zygentoma) and the winged of muscles allowing flapping movement remains insects for several characters related to the wings, unclear. In contrast, the Limb branch hypothesis e.g. thoracic muscles, muscles related to flight presumes that wings evolved in aquatic habitats. apparatus. However, recent head structure Based on the fossil record, winged insects analyses revived the Palaeoptera hypothesis – a probably had a terrestrial origin and the different sister-group relationship between dragonflies and adaptions of aquatic nymphs seen in basal winged mayflies (Odonata+Ephemeroptera, Neoptera) [7]. insects indicate a secondarily independent In contrast, establishing a sound phylogenetic colonization of aquatic habitats [4]. Paleontological hypothesis for the evolution of flying insects based evidence is limited in terms of addressing the on molecular data sets is problematic even in the question of morphological and functional origin of phylogenomic era, resulting in striking dis- insect wings. No fossil record is known with agreements; see [8, 9] and references therein. intermediate winged stages and the oldest Additionally, the elucidation of the origin of wing presumably winged insect fossil is 45 million years folding – another evolutionary novelty with critical (My) younger than primarily wingless ancestors importance for insect radiation – remains (411.5 My) [5]. However, expression studies of challenging. Within Neoptera three infraclasses three wing-regulatory genes in a mayfly have been proposed based on morphological (Ephemeroptera) and a wingless bristletail insect characters: Polyneoptera (lower neopteran insects; (Archaeognatha) proposed a combinatorial grasshoppers, mantids, earwigs, stoneflies, etc.), developmental model for wing origin where the Paraneoptera (thrips, lice, bugs in the wide sense) thoracic tergal edge (paranotal lobe) is the putative and Holometabola (insects undergoing complete source of wing margin formation [6]. metamorphosis; flies, beetles, butterflies, etc.). While the monophyly of the latter two is well Early evolution of winged insects established based on morphological and molecular Knowledge of the evolution of the earliest analyses, evidence for monophyletic Polyneoptera pterygote lineages is critical to understanding what has only slightly increased in the past. Analyses of ancestral structure wings evolved from. The wing base structures [10] and large transcriptomic traditional view separating Pterygota into two data sets [8, 11] consistently provide support for monophyletic clades – Palaeoptera and Neoptera, this grouping, while e.g. rRNA gene analyses, still based on the ability or inability of wing folding – favor a paraphyletic lower neopteran group as the has been disproved based on different sets of common ancestor of wing folding in Neoptera. The morphological characters. Indeed, morphological “ancient rapid radiation” [12] of the insects, their characters have provided strong support for all mega-diversity [13] and the preserved ancient three possible branching patterns between the two characters in some taxa [14] might all be extant palaeopterous orders – Ephemeroptera contributing factors which complicate a robust (mayflies) and Odonata (dragon- and damselflies) resolution of insect relationships and consequently (both unable to fold wings over abdomen) – and hamper the elucidation of insect wing evolution. Please note: this contribution was not peer-reviewed. 2 Acknowledgements This work was supported by a fellowship within the Postdoc-Program of the German Academic Exchange Service (DAAD). Special thanks go also to Rob DeSalle, my supervisor, who carefully checked this article. References 1. Jockusch EL, Ober KA. 2004. Hypothesis testing incongruence using new transcriptomic data. in evolutionary developmental biology: A case study Genome Biol Evol 4: 1295-309. from insect wings. J Hered 95: 382-96. 9. Simon S, Strauss S, von Haeseler A, Hadrys H. 2. Hamilton KGA. 1971. The insect wing, Pt. 1. 2009. A phylogenomic approach to resolve the Origin and development of wings from notal lobes. J basal pterygote divergence. Mol Biol Evol 26: 2719- Kansas Entomol Soc 44: 421-33. 30. 3. Averof M, Cohen SM. 1997. Evolutionary origin of 10. Yoshizawa K. 2011. Monophyletic Polyneoptera insect wings from ancestral gills. Nature 385: 627- recovered by wing base structure. Syst Entomol 36: 30. 377-94. 4. Grimaldi DA, Engel MS. 2005. Evolution of the 11. Letsch HO, Meusemann K, Wipfler B, Schutte K, Insects New York: Cambridge University Press. et al. 2012. Insect phylogenomics: results, problems 5. Garrouste R, Clement G, Nel P, Engel MS, et al. and the impact of matrix composition. Proc Biol Sci 2012. A complete insect from the Late Devonian 279: 3282-90. period. Nature 488: 82-5. 12. Whitfield JB, Kjer KM. 2008. Ancient rapid 6. Niwa N, Akimoto-Kato A, Niimi T, Tojo K, et al. radiations of insects: challenges for phylogenetic 2010. Evolutionary origin of the insect wing via analysis. Annu Rev Entomol 53: 449-72. integration of two developmental modules. Evol Dev 13. Yeates DK, Cameron SL, Trautwein M. 2012. A 12: 168-76. view from the edge of the forest: recent progress in 7. Blanke A, Wipfler B, Letsch H, Koch M, et al. understanding the relationships of the insect orders. 2012. Revival of Palaeoptera—head characters Aust J Entomol 51: 79-87. support a monophyletic origin of Odonata and 14. Willmann R. 2004. Phylogenetic Relationships and Ephemeroptera (Insecta). Cladistics 28: 560-81. Evolution of Insects. In Cracraft J, Donoghue MJ, 8. Simon S, Narechania A, Desalle R, Hadrys H. eds. Assembling the Tree of Life: Oxford University 2012. Insect phylogenomics: exploring the source of Press. p 330-44. .
Load more

Recommended publications
  • The Mitochondrial Genomes of Palaeopteran Insects and Insights
    www.nature.com/scientificreports OPEN The mitochondrial genomes of palaeopteran insects and insights into the early insect relationships Nan Song1*, Xinxin Li1, Xinming Yin1, Xinghao Li1, Jian Yin2 & Pengliang Pan2 Phylogenetic relationships of basal insects remain a matter of discussion. In particular, the relationships among Ephemeroptera, Odonata and Neoptera are the focus of debate. In this study, we used a next-generation sequencing approach to reconstruct new mitochondrial genomes (mitogenomes) from 18 species of basal insects, including six representatives of Ephemeroptera and 11 of Odonata, plus one species belonging to Zygentoma. We then compared the structures of the newly sequenced mitogenomes. A tRNA gene cluster of IMQM was found in three ephemeropteran species, which may serve as a potential synapomorphy for the family Heptageniidae. Combined with published insect mitogenome sequences, we constructed a data matrix with all 37 mitochondrial genes of 85 taxa, which had a sampling concentrating on the palaeopteran lineages. Phylogenetic analyses were performed based on various data coding schemes, using maximum likelihood and Bayesian inferences under diferent models of sequence evolution. Our results generally recovered Zygentoma as a monophyletic group, which formed a sister group to Pterygota. This confrmed the relatively primitive position of Zygentoma to Ephemeroptera, Odonata and Neoptera. Analyses using site-heterogeneous CAT-GTR model strongly supported the Palaeoptera clade, with the monophyletic Ephemeroptera being sister to the monophyletic Odonata. In addition, a sister group relationship between Palaeoptera and Neoptera was supported by the current mitogenomic data. Te acquisition of wings and of ability of fight contribute to the success of insects in the planet.
    [Show full text]
  • A New Insect Trackway from the Upper Jurassic—Lower Cretaceous Eolian Sandstones of São Paulo State, Brazil: Implications for Reconstructing Desert Paleoecology
    A new insect trackway from the Upper Jurassic—Lower Cretaceous eolian sandstones of São Paulo State, Brazil: implications for reconstructing desert paleoecology Bernardo de C.P. e M. Peixoto1,2, M. Gabriela Mángano3, Nicholas J. Minter4, Luciana Bueno dos Reis Fernandes1 and Marcelo Adorna Fernandes1,2 1 Laboratório de Paleoicnologia e Paleoecologia, Departamento de Ecologia e Biologia Evolutiva, Universidade Federal de São Carlos (UFSCar), São Carlos, São Paulo, Brazil 2 Programa de Pós Graduacão¸ em Ecologia e Recursos Naturais, Centro de Ciências Biológicas e da Saúde, Universidade Federal de São Carlos (UFSCar), São Carlos, São Paulo, Brazil 3 Department of Geological Sciences, University of Saskatchewan, Saskatoon, Saskatchewan, Canada 4 School of the Environment, Geography, and Geosciences, University of Portsmouth, Portsmouth, Hampshire, United Kingdom ABSTRACT The new ichnospecies Paleohelcura araraquarensis isp. nov. is described from the Upper Jurassic-Lower Cretaceous Botucatu Formation of Brazil. This formation records a gigantic eolian sand sea (erg), formed under an arid climate in the south-central part of Gondwana. This trackway is composed of two track rows, whose internal width is less than one-quarter of the external width, with alternating to staggered series, consisting of three elliptical tracks that can vary from slightly elongated to tapered or circular. The trackways were found in yellowish/reddish sandstone in a quarry in the Araraquara municipality, São Paulo State. Comparisons with neoichnological studies and morphological inferences indicate that the producer of Paleohelcura araraquarensis isp. nov. was most likely a pterygote insect, and so could have fulfilled one of the Submitted 6 November 2019 ecological roles that different species of this group are capable of performing in dune Accepted 10 March 2020 deserts.
    [Show full text]
  • Fossil Record of Stem Groups Employed In
    www.nature.com/scientificreports OPEN Fossil record of stem groups employed in evaluating the chronogram of insects (Arthropoda: Received: 07 April 2016 Accepted: 16 November 2016 Hexapoda) Published: 13 December 2016 Yan-hui Wang1,2,*, Michael S. Engel3,*, José A. Rafael4,*, Hao-yang Wu2, Dávid Rédei2, Qiang Xie2, Gang Wang1, Xiao-guang Liu1 & Wen-jun Bu2 Insecta s. str. (=Ectognatha), comprise the largest and most diversified group of living organisms, accounting for roughly half of the biodiversity on Earth. Understanding insect relationships and the specific time intervals for their episodes of radiation and extinction are critical to any comprehensive perspective on evolutionary events. Although some deeper nodes have been resolved congruently, the complete evolution of insects has remained obscure due to the lack of direct fossil evidence. Besides, various evolutionary phases of insects and the corresponding driving forces of diversification remain to be recognized. In this study, a comprehensive sample of all insect orders was used to reconstruct their phylogenetic relationships and estimate deep divergences. The phylogenetic relationships of insect orders were congruently recovered by Bayesian inference and maximum likelihood analyses. A complete timescale of divergences based on an uncorrelated log-normal relaxed clock model was established among all lineages of winged insects. The inferred timescale for various nodes are congruent with major historical events including the increase of atmospheric oxygen in the Late Silurian and earliest Devonian, the radiation of vascular plants in the Devonian, and with the available fossil record of the stem groups to various insect lineages in the Devonian and Carboniferous. Over half of all described living species are insects, and they dominate all terrestrial ecosystems1.
    [Show full text]
  • The Homology of Wing Base Sclerites and Flight Muscles In
    Arthropod Structure & Development 36 (2007) 253e269 www.elsevier.com/locate/asd The homology of wing base sclerites and flight muscles in Ephemeroptera and Neoptera and the morphology of the pterothorax of Habroleptoides confusa (Insecta: Ephemeroptera: Leptophlebiidae) Jana Willkommen*, Thomas Ho¨rnschemeyer1 Blumenbach-Institut fu¨r Zoologie & Anthropologie, Abt. Morphologie & Systematik, Berliner Str. 28, D-37073 Go¨ttingen, Germany Received 9 November 2006; accepted 11 January 2007 Abstract The ability to fly is the decisive factor for the evolutionary success of winged insects (Pterygota). Despite this, very little is known about the ground-pattern and evolution of the functionally very important wing base. Here we use the Ephemeroptera, usually regarded as the most ancient flying insects, as a model for the analysis of the flight musculature and the sclerites of the wing base. Morphology and anatomy of the pterothorax of 13 species of Ephemeroptera and five species of Plecoptera were examined and a detailed description of Habroleptoides confusa (Ephemero- ptera: Leptophlebiidae) is given. A new homology of the wing base sclerites in Ephemeroptera is proposed. The wing base of Ephemeroptera possesses three axillary sclerites that are homologous to the first axillary, the second axillary and the third axillary of Neoptera. For example, the third axillary possesses the axillary-pleural muscle that mostly is considered as a characteristic feature of the Neoptera. Many of the muscles and sclerites of the flight system of the Ephemeroptera and Neoptera can be readily homologised. In fact, there are indications that a foldable wing base may be a ground plan feature of pterygote insects and that the non-foldable wing base of the Ephemeroptera is a derived state.
    [Show full text]
  • Burmese Amber Taxa
    Burmese (Myanmar) amber taxa, on-line supplement v.2021.1 Andrew J. Ross 21/06/2021 Principal Curator of Palaeobiology Department of Natural Sciences National Museums Scotland Chambers St. Edinburgh EH1 1JF E-mail: [email protected] Dr Andrew Ross | National Museums Scotland (nms.ac.uk) This taxonomic list is a supplement to Ross (2021) and follows the same format. It includes taxa described or recorded from the beginning of January 2021 up to the end of May 2021, plus 3 species that were named in 2020 which were missed. Please note that only higher taxa that include new taxa or changed/corrected records are listed below. The list is until the end of May, however some papers published in June are listed in the ‘in press’ section at the end, but taxa from these are not yet included in the checklist. As per the previous on-line checklists, in the bibliography page numbers have been added (in blue) to those papers that were published on-line previously without page numbers. New additions or changes to the previously published list and supplements are marked in blue, corrections are marked in red. In Ross (2021) new species of spider from Wunderlich & Müller (2020) were listed as being authored by both authors because there was no indication next to the new name to indicate otherwise, however in the introduction it was indicated that the author of the new taxa was Wunderlich only. Where there have been subsequent taxonomic changes to any of these species the authorship has been corrected below.
    [Show full text]
  • Columbia County Since 1881 Leap Day Coronavirus Strikes Oregon Babies Celebrate
    ‘Spring Equipment ahead’ fire at area Sunday log yard Page A4 March 8 Page A3 Wednesday, March 4, 2020 $1 TheThe ChronicleChronicle thechronicleonline.com Serving Columbia County since 1881 Leap Day Coronavirus Strikes Oregon babies celebrate CHRISTINE MENGES [email protected] This past Saturday, the world had a Leap Day, or an extra day added to February, Feb. 29. Most years, February ends on the 28th. But once every four years, the month gains an extra day. The reason why has to do with how the earth revolves around the sun. While people tend to think of years as being 365 days long, in reality it takes the earth 365 days, five hours, 48 minutes and 46 seconds to make a full revolution. If calendars didn’t account for those extra five-ish hours, the seasons would eventually go out of whack. That’s why, to account for the discrep- ancy, an extra day is added (more or Oregon cases increasing School districts take action Columbia County response less) every four years. For most people, the extra day JEREMY C. RUARK JEREMY C. RUARK CHRISTINE MENGES is just one more day in the calendar [email protected] [email protected] [email protected] year. For those born on the date, it’s a Oregon health officials have identified School officials in Columbia County The Chronicle sat down with Michael bigger deal. a third presumptive positive case of CO- are taking preventive steps to ensure Paul, the Public Health Administrator for While most people have a 1 out VID-19 among state residents.
    [Show full text]
  • Is Ellipura Monophyletic? a Combined Analysis of Basal Hexapod
    ARTICLE IN PRESS Organisms, Diversity & Evolution 4 (2004) 319–340 www.elsevier.de/ode Is Ellipura monophyletic? A combined analysis of basal hexapod relationships with emphasis on the origin of insects Gonzalo Giribeta,Ã, Gregory D.Edgecombe b, James M.Carpenter c, Cyrille A.D’Haese d, Ward C.Wheeler c aDepartment of Organismic and Evolutionary Biology, Museum of Comparative Zoology, Harvard University, 16 Divinity Avenue, Cambridge, MA 02138, USA bAustralian Museum, 6 College Street, Sydney, New South Wales 2010, Australia cDivision of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, USA dFRE 2695 CNRS, De´partement Syste´matique et Evolution, Muse´um National d’Histoire Naturelle, 45 rue Buffon, F-75005 Paris, France Received 27 February 2004; accepted 18 May 2004 Abstract Hexapoda includes 33 commonly recognized orders, most of them insects.Ongoing controversy concerns the grouping of Protura and Collembola as a taxon Ellipura, the monophyly of Diplura, a single or multiple origins of entognathy, and the monophyly or paraphyly of the silverfish (Lepidotrichidae and Zygentoma s.s.) with respect to other dicondylous insects.Here we analyze relationships among basal hexapod orders via a cladistic analysis of sequence data for five molecular markers and 189 morphological characters in a simultaneous analysis framework using myriapod and crustacean outgroups.Using a sensitivity analysis approach and testing for stability, the most congruent parameters resolve Tricholepidion as sister group to the remaining Dicondylia, whereas most suboptimal parameter sets group Tricholepidion with Zygentoma.Stable hypotheses include the monophyly of Diplura, and a sister group relationship between Diplura and Protura, contradicting the Ellipura hypothesis.Hexapod monophyly is contradicted by an alliance between Collembola, Crustacea and Ectognatha (i.e., exclusive of Diplura and Protura) in molecular and combined analyses.
    [Show full text]
  • Stable Structural Color Patterns Displayed on Transparent Insect Wings
    Stable structural color patterns displayed on transparent insect wings Ekaterina Shevtsovaa,1, Christer Hanssona,b,1, Daniel H. Janzenc,1, and Jostein Kjærandsend,1 aDepartment of Biology, Lund University, Sölvegatan 35, SE-22362 Lund, Sweden; bScientific Associate of the Entomology Department, Natural History Museum, London SW7 5BD, United Kingdom; cDepartment of Biology, University of Pennsylvania, Philadelphia, PA 19104-6018; and dDepartment of Biology, Museum of Zoology, Lund University, Helgonavägen 3, SE-22362 Lund, Sweden Contributed by Daniel H. Janzen, November 24, 2010 (sent for review October 5, 2010) Color patterns play central roles in the behavior of insects, and are and F). In laboratory conditions most wings are studied against a important traits for taxonomic studies. Here we report striking and white background (Fig. 1 G, H, and J), or the wings are embedded stable structural color patterns—wing interference patterns (WIPs) in a medium with a refractive index close to that of chitin (e.g., —in the transparent wings of small Hymenoptera and Diptera, ref. 19). In both cases the color reflections will be faint or in- patterns that have been largely overlooked by biologists. These ex- visible. tremely thin wings reflect vivid color patterns caused by thin film Insects are an exceedingly diverse and ancient group and interference. The visibility of these patterns is affected by the way their signal-receiver architecture of thin membranous wings the insects display their wings against various backgrounds with and color vision was apparently in place before their huge radia- different light properties. The specific color sequence displayed tion (20–22). The evolution of functional wings (Pterygota) that lacks pure red and matches the color vision of most insects, strongly can be freely operated in multidirections (Neoptera), coupled suggesting that the biological significance of WIPs lies in visual with small body size, has long been viewed as associated with their signaling.
    [Show full text]
  • Arthropods of Elm Fork Preserve
    Arthropods of Elm Fork Preserve Arthropods are characterized by having jointed limbs and exoskeletons. They include a diverse assortment of creatures: Insects, spiders, crustaceans (crayfish, crabs, pill bugs), centipedes and millipedes among others. Column Headings Scientific Name: The phenomenal diversity of arthropods, creates numerous difficulties in the determination of species. Positive identification is often achieved only by specialists using obscure monographs to ‘key out’ a species by examining microscopic differences in anatomy. For our purposes in this survey of the fauna, classification at a lower level of resolution still yields valuable information. For instance, knowing that ant lions belong to the Family, Myrmeleontidae, allows us to quickly look them up on the Internet and be confident we are not being fooled by a common name that may also apply to some other, unrelated something. With the Family name firmly in hand, we may explore the natural history of ant lions without needing to know exactly which species we are viewing. In some instances identification is only readily available at an even higher ranking such as Class. Millipedes are in the Class Diplopoda. There are many Orders (O) of millipedes and they are not easily differentiated so this entry is best left at the rank of Class. A great deal of taxonomic reorganization has been occurring lately with advances in DNA analysis pointing out underlying connections and differences that were previously unrealized. For this reason, all other rankings aside from Family, Genus and Species have been omitted from the interior of the tables since many of these ranks are in a state of flux.
    [Show full text]
  • Consequences of Evolutionary Transitions in Changing Photic Environments
    bs_bs_banner Austral Entomology (2017) 56,23–46 Review Consequences of evolutionary transitions in changing photic environments Simon M Tierney,1* Markus Friedrich,2,3 William F Humphreys,1,4,5 Therésa M Jones,6 Eric J Warrant7 and William T Wcislo8 1School of Biological Sciences, The University of Adelaide, North Terrace, Adelaide, SA 5005, Australia. 2Department of Biological Sciences, Wayne State University, 5047 Gullen Mall, Detroit, MI 48202, USA. 3Department of Anatomy and Cell Biology, Wayne State University, School of Medicine, 540 East Canfield Avenue, Detroit, MI 48201, USA. 4Terrestrial Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, WA 6986, Australia. 5School of Animal Biology, University of Western Australia, Nedlands, WA 6907, Australia. 6Department of Zoology, The University of Melbourne, Melbourne, Vic. 3010, Australia. 7Department of Biology, Lund University, Sölvegatan 35, S-22362 Lund, Sweden. 8Smithsonian Tropical Research Institute, PO Box 0843-03092, Balboa, Ancón, Republic of Panamá. Abstract Light represents one of the most reliable environmental cues in the biological world. In this review we focus on the evolutionary consequences to changes in organismal photic environments, with a specific focus on the class Insecta. Particular emphasis is placed on transitional forms that can be used to track the evolution from (1) diurnal to nocturnal (dim-light) or (2) surface to subterranean (aphotic) environments, as well as (3) the ecological encroachment of anthropomorphic light on nocturnal habitats (artificial light at night). We explore the influence of the light environment in an integrated manner, highlighting the connections between phenotypic adaptations (behaviour, morphology, neurology and endocrinology), molecular genetics and their combined influence on organismal fitness.
    [Show full text]
  • Volume 2, Chapter 12-5: Terrestrial Insects: Hemimetabola-Notoptera
    Glime, J. M. 2017. Terrestrial Insects: Hemimetabola – Notoptera and Psocoptera. Chapter 12-5. In: Glime, J. M. Bryophyte Ecology. 12-5-1 Volume 2. Interactions. Ebook sponsored by Michigan Technological University and the International Association of Bryologists. eBook last updated 19 July 2020 and available at <http://digitalcommons.mtu.edu/bryophyte-ecology2/>. CHAPTER 12-5 TERRESTRIAL INSECTS: HEMIMETABOLA – NOTOPTERA AND PSOCOPTERA TABLE OF CONTENTS NOTOPTERA .................................................................................................................................................. 12-5-2 Grylloblattodea – Ice Crawlers ................................................................................................................. 12-5-3 Grylloblattidae – Ice Crawlers ........................................................................................................... 12-5-3 Galloisiana ................................................................................................................................. 12-5-3 Grylloblatta ................................................................................................................................ 12-5-3 Grylloblattella ............................................................................................................................ 12-5-4 PSOCOPTERA – Booklice, Barklice, Barkflies .............................................................................................. 12-5-4 Summary .........................................................................................................................................................
    [Show full text]
  • Description of an Eyeless Species of the Ground Beetle Genus Trechus Clairville, 1806 (Coleoptera: Carabidae: Trechini)
    Zootaxa 4083 (3): 431–443 ISSN 1175-5326 (print edition) http://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2016 Magnolia Press ISSN 1175-5334 (online edition) http://doi.org/10.11646/zootaxa.4083.3.7 http://zoobank.org/urn:lsid:zoobank.org:pub:C999EBFD-4EAF-44E1-B7E9-95C9C63E556B Blind life in the Baltic amber forests: description of an eyeless species of the ground beetle genus Trechus Clairville, 1806 (Coleoptera: Carabidae: Trechini) JOACHIM SCHMIDT1, 2, HANNES HOFFMANN3 & PETER MICHALIK3 1University of Rostock, Institute of Biosciences, General and Systematic Zoology, Universitätsplatz 2, 18055 Rostock, Germany 2Lindenstraße 3a, 18211 Admannshagen, Germany. E-mail: [email protected] 3Zoological Institute and Museum, Ernst-Moritz-Arndt-University, Loitzer Str. 26, D-17489 Greifswald, Germany. E-mail: [email protected] Abstract The first eyeless beetle known from Baltic amber, Trechus eoanophthalmus sp. n., is described and imaged using light microscopy and X-ray micro-computed tomography. Based on external characters, the new species is most similar to spe- cies of the Palaearctic Trechus sensu stricto clade and seems to be closely related to the Baltic amber fossil T. balticus Schmidt & Faille, 2015. Due to the poor conservation of the internal parts of the body, no information on the genital char- acters can be provided. Therefore, the systematic position of this fossil within the megadiverse genus Trechus remains dubious. The occurrence of the blind and flightless T. eoanophthalmus sp. n. in the Baltic amber forests supports a previ- ous hypothesis that these forests were located in an area partly characterised by mountainous habitats with temperate cli- matic conditions.
    [Show full text]