DOCSLIB.ORG

Fossil Calibrations for the Arthropod Tree of Life

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
Fossil Calibrations for the Arthropod Tree of Life bioRxiv preprint doi: https://doi.org/10.1101/044859; this version posted June 10, 2016. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY 4.0 International license. FOSSIL CALIBRATIONS FOR THE ARTHROPOD TREE OF LIFE AUTHORS Joanna M. Wolfe1*, Allison C. Daley2,3, David A. Legg3, Gregory D. Edgecombe4 1 Department of Earth, Atmospheric & Planetary Sciences, Massachusetts Institute of Technology, Cambridge, MA 02139, USA 2 Department of Zoology, University of Oxford, South Parks Road, Oxford OX1 3PS, UK 3 Oxford University Museum of Natural History, Parks Road, Oxford OX1 3PZ, UK 4 Department of Earth Sciences, The Natural History Museum, Cromwell Road, London SW7 5BD, UK *Corresponding author: [email protected] ABSTRACT Fossil age data and molecular sequences are increasingly combined to establish a timescale for the Tree of Life. Arthropods, as the most species-rich and morphologically disparate animal phylum, have received substantial attention, particularly with regard to questions such as the timing of habitat shifts (e.g. terrestrialisation), genome evolution (e.g. gene family duplication and functional evolution), origins of novel characters and behaviours (e.g. wings and flight, venom, silk), biogeography, rate of diversification (e.g. Cambrian explosion, insect coevolution with angiosperms, evolution of crab body plans), and the evolution of arthropod microbiomes. We present herein a series of rigorously vetted calibration fossils for arthropod evolutionary history, taking into account recently published guidelines for best practice in fossil calibration. These are restricted to Palaeozoic and Mesozoic fossils, no deeper than ordinal taxonomic level, nonetheless resulting in 80 fossil calibrations for 102 clades. This work is especially timely owing to the rapid growth of molecular sequence data and the fact that many included fossils have been described within the last five years. This contribution provides a resource for systematists and other biologists interested in deep-time questions in arthropod evolution. KEYWORDS Arthropods; Fossils; Phylogeny; Divergence times ABBREVIATIONS AMNH, American Museum of Natural History; AMS, Australian Museum, Sydney; AUGD, University of Aberdeen; BGR, Bundesanstalt für Geowissenschaften und Rohstoffe, Berlin; BMNH, The Natural History Museum, London; CNU, Key Laboratory of Insect Evolutionary & Environmental Change, Capital Normal University, Beijing; DE, Ulster Museum, Belfast; ED, Ibaraki University, Mito, Japan; FMNH, Field Museum of Natural History; GMCB, Geological Museum of China, Beijing; GSC, Geological Survey of Canada; IRNSB, Institut Royal des Sciences Naturelles de Belgique, Brussels; KSU, Kent State University; Ld, Musée Fleury, Lodève, France; LWL, Landschaftsverband Westfalen- Lippe-Museum für Naturkunde, Münster; MACN, Museo Argentino de Ciencias Naturales, Buenos Aires; MBA, Museum für Naturkunde, Berlin; MCNA, Museo de Ciencias Naturales de Álava, Vitoria- Gasteiz, Álava, Spain; MCZ, Museum of Comparative Zoology, Harvard University; MGSB, Museo Geologico del Seminario de Barcelona; MN, Museu Nacional, Rio de Janeiro; MNHN, Muséum national d'Histoire naturelle, Paris; NHMUK, The Natural History Museum, London; NIGP, Nanjing Institute of Geology and Palaeontology; NMS, National Museum of Scotland; OUM, Oxford 1 bioRxiv preprint doi: https://doi.org/10.1101/044859; this version posted June 10, 2016. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY 4.0 International license. University Museum of Natural History; PBM, Palaöbotanik Münster; PIN, Paleontological Institute, Moscow; PRI, Paleontological Research Institution, Ithaca; ROM, Royal Ontario Museum; SAM, South Australian Museum, Adelaide; SM, Sedgwick Museum, University of Cambridge; SMNK, Staatliches Museum für Naturkunde, Karlsruhe; SMNS, Staatliches Museum für Naturkunde, Stuttgart; TsGM, F.N. Chernyshev Central Geologic Prospecting Research Museum, St. Petersburg; UB, University of Bonn; USNM, US National Museum of Natural History, Smithsonian Institution; UWGM, University of Wisconsin Geology Museum; YKLP, Yunnan Key Laboratory for Palaeobiology, Yunnan University; YPM, Yale Peabody Museum; ZPAL, Institute of Paleobiology, Polish Academy of Sciences, Warsaw. 1. Introduction Accurate and precise systematic placement and dating of fossils underpins most efforts to infer a chronology for the Tree of Life. Arthropods, as a whole or in part, have received considerable focus owing to their incredible morphological disparity, species richness, and (relative to much of the Tree of Life) excellent fossil record. A growing number of recent studies have constructed timetrees for arthropods as a whole or for major groups therein (e.g. Bellec and Rabet, 2016; Bond et al., 2014; Bracken-Grissom et al., 2014, 2013; Djernæs et al., 2015; Fernández et al., 2016, 2014; Fernández and Giribet, 2015; Garrison et al., 2016; Garwood et al., 2014; Giribet and Edgecombe, 2013; Herrera et al., 2015; Klopfstein et al., 2015; Legendre et al., 2015; Malm et al., 2013; McKenna et al., 2015; Misof et al., 2014; Oakley et al., 2013; Rehm et al., 2011; Schwentner et al., 2013; Song et al., 2015; Sun et al., 2015; Thomas et al., 2013; Tsang et al., 2014; Wahlberg et al., 2013; Wiegmann et al., 2011; Wood et al., 2013; Xu et al., 2015; Zhu et al., 2015). These studies vary in how well they have adhered to best practices for selecting calibration fossils, as many previous calibrations assume that fossil taxonomy accurately reflects phylogeny. Compounding the issue is the expansion of divergence time studies for a variety of comparative questions far beyond systematics and biogeography, including habitat shifts (Letsch et al., 2016; Lins et al., 2012; Rota-Stabelli et al., 2013a; Yang et al., 2013), genome evolution (Cao et al., 2013; Schwarz et al., 2014; Starrett et al., 2013; Wissler et al., 2013; Yuan et al., 2016), origins of novel characters and behaviours (Rainford et al., 2014; Sanggaard et al., 2014; Wheat and Wahlberg, 2013), evolution of parasites and disease (Ibarra-Cerdeña et al., 2014; Palopoli et al., 2014; Rees et al., 2014; Zhou et al., 2014), rate of diversification and its relationship to morphology and ecology (Lee et al., 2013; Wiens et al., 2015), coevolution (Kaltenpoth et al., 2014; Shelomi et al., 2016; Wilson et al., 2013), conservation (Owen et al., 2015), and the use of arthropods as a model for methodological development (O’Reilly et al., 2015; Ronquist et al., 2012; Warnock et al., 2012; Zhang et al., 2016). Recent consensus on best practices for calibration fossil selection requires reference to specific fossil specimen(s), phylogenetic or morphological evidence justifying placement of the fossil, and stratigraphic and/or absolute dating information for the fossil (Parham et al., 2012). The importance of accurate phylogenetic knowledge of calibration fossils is underscored by recent controversies in dating the evolution of insects, where arguments hinge on the classification of particular ‘roachoid’ fossils on the stem lineage of Dictyoptera, with resulting differences on the order of 100 Myr (Kjer et al., 2015; Tong et al., 2015). With the explosion of taxonomic sampling in molecular phylogenies due to improvements in sequencing technology, improving the coverage of fossil calibrations is equally important. Recommendations include, for example, including as many as one fossil per ten extant OTUs for precise ages, with a varied distribution across lineages and clade depth (Bracken-Grissom et al., 2014). As a response, we have compiled an atlas of 80 rigorously scrutinized calibration fossils for 102 key nodes in arthropod phylogeny. These represent four basal ecdysozoan and arthropod clades, 17 2 bioRxiv preprint doi: https://doi.org/10.1101/044859; this version posted June 10, 2016. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY 4.0 International license. chelicerates, 12 myriapods, 30 non-hexapod pancrustaceans, and 39 hexapod clades. Where possible, we favour clade topologies resulting from a phylogenetic analysis of the largest total dataset. If phylogenomic analysis of genomes or transcriptomes has been performed but conflicts with morphology, a strongly supported molecular result is presented (e.g. putative clades such as Oligostraca that do not yet have identified morphological autapomorphies). If, however, molecular phylogenies have been constructed with few genes (e.g. clades such as Peracarida) or with highly conflicting results (e.g. Arachnida), morphological results are given greater weight. Where relevant, we discuss clade names with respect to NCBI’s GenBank taxonomy (as recommended by the Fossil Calibrations Database: Polly et al., 2015), as this review is intended to be used by molecular biologists who are interested in dating the evolution of arthropod groups. As there are >1.2 million species of arthropods, our calibrations are limited to fossils from the Palaeozoic and Mesozoic. Many extant clades have their oldest fossils in Cenozoic ambers such as the Eocene Baltic amber but are predicted to be vastly older based
Load more

Recommended publications
  • Mechanoreceptors in Early Developmental Stages of the Pycnogonida
    UACE2019 - Conference Proceedings Mechanoreceptors in Early Developmental Stages of the Pycnogonida John A. Fornshell * a a U.S. National Museum of Natural History Department of Invertebrate Zoology Smithsonian Institution Washington, D.C. USA *Correspondence: [email protected]; Tel. (571) 426-2398 ABSTRACT Members of the phylum Arthropoda detect fluid flow and sound/particle vibrations using sensory organs called sensilla. These sensilla detect sound/particle vibrations in the boundary layer. In the present study, archived specimens from the United States National Museum of Natural History were examined in an effort to extend our knowledge of the presence of sensilla on the early post hatching developmental stages, first and second instars, of pycnogonids. In the work presented here we look at three families, four genera and ten species of early post hatching developmental stages of sea spiders. They are Family Ammotheidae, Achelia cuneatis Child, 1999, Ammothea allopodes Fry and Hedgpeth, 1969, Ammothea carolinensis Leach 1814, Ammothea clausi Pfeffer, 1889, Ammothea striata (Möbius, 1902), Family Nymphonidae, Nymphon grossipes (Fabricius, 1780), N. australe Hodgson, 1902, N. charcoti Bouvier, 1911, N. Tenellum (Sars, 1888) and Pycnogonidae, Pentapycnon charcoti Bouvier, 1910. Electron micrograph images of these species were used to identify and describe the sensilla present. Most body organs such as mouthparts, the eye tubercle, appendages and spines are proportionally much smaller in the early post hatching developmental stages compared to their size in the adults, while the sensilla are comparable in size and shape to those found on the adults. In the first instar of Pentapycnon charcoti sensilla are present, but not in the adult.
    [Show full text]
  • Insecta: Phasmatodea) and Their Phylogeny
    insects Article Three Complete Mitochondrial Genomes of Orestes guangxiensis, Peruphasma schultei, and Phryganistria guangxiensis (Insecta: Phasmatodea) and Their Phylogeny Ke-Ke Xu 1, Qing-Ping Chen 1, Sam Pedro Galilee Ayivi 1 , Jia-Yin Guan 1, Kenneth B. Storey 2, Dan-Na Yu 1,3 and Jia-Yong Zhang 1,3,* 1 College of Chemistry and Life Science, Zhejiang Normal University, Jinhua 321004, China; [email protected] (K.-K.X.); [email protected] (Q.-P.C.); [email protected] (S.P.G.A.); [email protected] (J.-Y.G.); [email protected] (D.-N.Y.) 2 Department of Biology, Carleton University, Ottawa, ON K1S 5B6, Canada; [email protected] 3 Key Lab of Wildlife Biotechnology, Conservation and Utilization of Zhejiang Province, Zhejiang Normal University, Jinhua 321004, China * Correspondence: [email protected] or [email protected] Simple Summary: Twenty-seven complete mitochondrial genomes of Phasmatodea have been published in the NCBI. To shed light on the intra-ordinal and inter-ordinal relationships among Phas- matodea, more mitochondrial genomes of stick insects are used to explore mitogenome structures and clarify the disputes regarding the phylogenetic relationships among Phasmatodea. We sequence and annotate the first acquired complete mitochondrial genome from the family Pseudophasmati- dae (Peruphasma schultei), the first reported mitochondrial genome from the genus Phryganistria Citation: Xu, K.-K.; Chen, Q.-P.; Ayivi, of Phasmatidae (P. guangxiensis), and the complete mitochondrial genome of Orestes guangxiensis S.P.G.; Guan, J.-Y.; Storey, K.B.; Yu, belonging to the family Heteropterygidae. We analyze the gene composition and the structure D.-N.; Zhang, J.-Y.
    [Show full text]
  • Research Report110
    ~ ~ WISCONSIN DEPARTMENT OF NATURAL RESOURCES A Survey of Rare and Endangered Mayflies of Selected RESEARCH Rivers of Wisconsin by Richard A. Lillie REPORT110 Bureau of Research, Monona December 1995 ~ Abstract The mayfly fauna of 25 rivers and streams in Wisconsin were surveyed during 1991-93 to document the temporal and spatial occurrence patterns of two state endangered mayflies, Acantha­ metropus pecatonica and Anepeorus simplex. Both species are candidates under review for addition to the federal List of Endang­ ered and Threatened Wildlife. Based on previous records of occur­ rence in Wisconsin, sampling was conducted during the period May-July using a combination of sampling methods, including dredges, air-lift pumps, kick-nets, and hand-picking of substrates. No specimens of Anepeorus simplex were collected. Three specimens (nymphs or larvae) of Acanthametropus pecatonica were found in the Black River, one nymph was collected from the lower Wisconsin River, and a partial exuviae was collected from the Chippewa River. Homoeoneuria ammophila was recorded from Wisconsin waters for the first time from the Black River and Sugar River. New site distribution records for the following Wiscon­ sin special concern species include: Macdunnoa persimplex, Metretopus borealis, Paracloeodes minutus, Parameletus chelifer, Pentagenia vittigera, Cercobrachys sp., and Pseudiron centra/is. Collection of many of the aforementioned species from large rivers appears to be dependent upon sampling sand-bottomed substrates at frequent intervals, as several species were relatively abundant during only very short time spans. Most species were associated with sand substrates in water < 2 m deep. Acantha­ metropus pecatonica and Anepeorus simplex should continue to be listed as endangered for state purposes and receive a biological rarity ranking of critically imperiled (S1 ranking), and both species should be considered as candidates proposed for listing as endangered or threatened as defined by the Endangered Species Act.
    [Show full text]
  • Biochemical Divergence Between Cavernicolous and Marine
    The position of crustaceans within Arthropoda - Evidence from nine molecular loci and morphology GONZALO GIRIBET', STEFAN RICHTER2, GREGORY D. EDGECOMBE3 & WARD C. WHEELER4 Department of Organismic and Evolutionary- Biology, Museum of Comparative Zoology; Harvard University, Cambridge, Massachusetts, U.S.A. ' Friedrich-Schiller-UniversitdtJena, Instituifiir Spezielte Zoologie und Evolutionsbiologie, Jena, Germany 3Australian Museum, Sydney, NSW, Australia Division of Invertebrate Zoology, American Museum of Natural History, New York, U.S.A. ABSTRACT The monophyly of Crustacea, relationships of crustaceans to other arthropods, and internal phylogeny of Crustacea are appraised via parsimony analysis in a total evidence frame­ work. Data include sequences from three nuclear ribosomal genes, four nuclear coding genes, and two mitochondrial genes, together with 352 characters from external morphol­ ogy, internal anatomy, development, and mitochondrial gene order. Subjecting the com­ bined data set to 20 different parameter sets for variable gap and transversion costs, crusta­ ceans group with hexapods in Tetraconata across nearly all explored parameter space, and are members of a monophyletic Mandibulata across much of the parameter space. Crustacea is non-monophyletic at low indel costs, but monophyly is favored at higher indel costs, at which morphology exerts a greater influence. The most stable higher-level crusta­ cean groupings are Malacostraca, Branchiopoda, Branchiura + Pentastomida, and an ostracod-cirripede group. For combined data, the Thoracopoda and Maxillopoda concepts are unsupported, and Entomostraca is only retrieved under parameter sets of low congruence. Most of the current disagreement over deep divisions in Arthropoda (e.g., Mandibulata versus Paradoxopoda or Cormogonida versus Chelicerata) can be viewed as uncertainty regarding the position of the root in the arthropod cladogram rather than as fundamental topological disagreement as supported in earlier studies (e.g., Schizoramia versus Mandibulata or Atelocerata versus Tetraconata).
    [Show full text]
  • Arachnida, Solifugae) with Special Focus on Functional Analyses and Phylogenetic Interpretations
    HISTOLOGY AND ULTRASTRUCTURE OF SOLIFUGES Comparative studies of organ systems of solifuges (Arachnida, Solifugae) with special focus on functional analyses and phylogenetic interpretations HISTOLOGIE UND ULTRASTRUKTUR DER SOLIFUGEN Vergleichende Studien an Organsystemen der Solifugen (Arachnida, Solifugae) mit Schwerpunkt auf funktionellen Analysen und phylogenetischen Interpretationen I N A U G U R A L D I S S E R T A T I O N zur Erlangung des akademischen Grades doctor rerum naturalium (Dr. rer. nat.) an der Mathematisch-Naturwissenschaftlichen Fakultät der Ernst-Moritz-Arndt-Universität Greifswald vorgelegt von Anja Elisabeth Klann geboren am 28.November 1976 in Bremen Greifswald, den 04.06.2009 Dekan ........................................................................................................Prof. Dr. Klaus Fesser Prof. Dr. Dr. h.c. Gerd Alberti Erster Gutachter .......................................................................................... Zweiter Gutachter ........................................................................................Prof. Dr. Romano Dallai Tag der Promotion ........................................................................................15.09.2009 Content Summary ..........................................................................................1 Zusammenfassung ..........................................................................5 Acknowledgments ..........................................................................9 1. Introduction ............................................................................
    [Show full text]
  • BÖCEKLERİN SINIFLANDIRILMASI (Takım Düzeyinde)
    BÖCEKLERİN SINIFLANDIRILMASI (TAKIM DÜZEYİNDE) GÖKHAN AYDIN 2016 Editör : Gökhan AYDIN Dizgi : Ziya ÖNCÜ ISBN : 978-605-87432-3-6 Böceklerin Sınıflandırılması isimli eğitim amaçlı hazırlanan bilgisayar programı için lütfen aşağıda verilen linki tıklayarak programı ücretsiz olarak bilgisayarınıza yükleyin. http://atabeymyo.sdu.edu.tr/assets/uploads/sites/76/files/siniflama-05102016.exe Eğitim Amaçlı Bilgisayar Programı ISBN: 978-605-87432-2-9 İçindekiler İçindekiler i Önsöz vi 1. Protura - Coneheads 1 1.1 Özellikleri 1 1.2 Ekonomik Önemi 2 1.3 Bunları Biliyor musunuz? 2 2. Collembola - Springtails 3 2.1 Özellikleri 3 2.2 Ekonomik Önemi 4 2.3 Bunları Biliyor musunuz? 4 3. Thysanura - Silverfish 6 3.1 Özellikleri 6 3.2 Ekonomik Önemi 7 3.3 Bunları Biliyor musunuz? 7 4. Microcoryphia - Bristletails 8 4.1 Özellikleri 8 4.2 Ekonomik Önemi 9 5. Diplura 10 5.1 Özellikleri 10 5.2 Ekonomik Önemi 10 5.3 Bunları Biliyor musunuz? 11 6. Plocoptera – Stoneflies 12 6.1 Özellikleri 12 6.2 Ekonomik Önemi 12 6.3 Bunları Biliyor musunuz? 13 7. Embioptera - webspinners 14 7.1 Özellikleri 15 7.2 Ekonomik Önemi 15 7.3 Bunları Biliyor musunuz? 15 8. Orthoptera–Grasshoppers, Crickets 16 8.1 Özellikleri 16 8.2 Ekonomik Önemi 16 8.3 Bunları Biliyor musunuz? 17 i 9. Phasmida - Walkingsticks 20 9.1 Özellikleri 20 9.2 Ekonomik Önemi 21 9.3 Bunları Biliyor musunuz? 21 10. Dermaptera - Earwigs 23 10.1 Özellikleri 23 10.2 Ekonomik Önemi 24 10.3 Bunları Biliyor musunuz? 24 11. Zoraptera 25 11.1 Özellikleri 25 11.2 Ekonomik Önemi 25 11.3 Bunları Biliyor musunuz? 26 12.
    [Show full text]
  • Phylogenomic Resolution of Sea Spider Diversification Through Integration Of
    bioRxiv preprint doi: https://doi.org/10.1101/2020.01.31.929612; this version posted February 2, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. Phylogenomic resolution of sea spider diversification through integration of multiple data classes 1Jesús A. Ballesteros†, 1Emily V.W. Setton†, 1Carlos E. Santibáñez López†, 2Claudia P. Arango, 3Georg Brenneis, 4Saskia Brix, 5Esperanza Cano-Sánchez, 6Merai Dandouch, 6Geoffrey F. Dilly, 7Marc P. Eleaume, 1Guilherme Gainett, 8Cyril Gallut, 6Sean McAtee, 6Lauren McIntyre, 9Amy L. Moran, 6Randy Moran, 5Pablo J. López-González, 10Gerhard Scholtz, 6Clay Williamson, 11H. Arthur Woods, 12Ward C. Wheeler, 1Prashant P. Sharma* 1 Department of Integrative Biology, University of Wisconsin–Madison, Madison, WI, USA 2 Queensland Museum, Biodiversity Program, Brisbane, Australia 3 Zoologisches Institut und Museum, Cytologie und Evolutionsbiologie, Universität Greifswald, Greifswald, Germany 4 Senckenberg am Meer, German Centre for Marine Biodiversity Research (DZMB), c/o Biocenter Grindel (CeNak), Martin-Luther-King-Platz 3, Hamburg, Germany 5 Biodiversidad y Ecología Acuática, Departamento de Zoología, Facultad de Biología, Universidad de Sevilla, Sevilla, Spain 6 Department of Biology, California State University-Channel Islands, Camarillo, CA, USA 7 Départment Milieux et Peuplements Aquatiques, Muséum national d’Histoire naturelle, Paris, France 8 Institut de Systématique, Emvolution, Biodiversité (ISYEB), Sorbonne Université, CNRS, Concarneau, France 9 Department of Biology, University of Hawai’i at Mānoa, Honolulu, HI, USA Page 1 of 31 bioRxiv preprint doi: https://doi.org/10.1101/2020.01.31.929612; this version posted February 2, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder.
    [Show full text]
  • Phasmid Studies
    ISSN 0966-0011 PHASMID STUDIES. volume 13, numbers 1 & 2. September 2005. Editor: P.E. Bragg. Published by the Phasmid Study Group. Phasmid Studies ISSN 0966-0011 volume 13, numbers I & 2. Contents Phasmids from Sabah Robert Bradburne I A reassessment of some Bornean Lonchodinae and Aschiphasmalidae, with some lectotype designations, new synonyms, and the description of (WO new species P.E. Bragg ................ ........ .. 11 Hap/opus Burmeisler, 1838, replacement name for Aplopus Gray, 1835 (Phasmalodea). Oliver Zornpro ... .. 30 A new species of the genus Baculofraclum. the first record of the genus from Borneo. P.E. Bragg .. ............................. .. 31 Reviews and Abstracts Phasmid Abstracts 38 Cover illustration: Female Parafoxopsis kQrySll!.~ Gilmher, 1932 by r.E. Bragg. Br.dburn, R. (2005) Phasmid Studies, 13(1&2): \-10. - Phasmids from Sabah Robert Bradburne, 26 Royal Avenue, Tonbridge, Kent, TN9 208, UK. Abstract This paper describes a trip (Q six locations in Sabah, Borneo, during October 2003. A 101:11 of around 20 species of stick insects were found al four of these locations, including an undescribed species found at 3300m on Moum Kinabalu. The most commonly encountered species in the lowland forest were Lonchodes spp., Haaniella spp., and Asceles margarilatus. Key words Phasmida, Borneo, Sabah, Sukau, Kinabalu, Danum Valley, Haaniella, Asceles, Prosemoria, Necroscia, Presbistus, Carausius, PhellQcephoms, Dinophasmo. Introduction In October 2003 I travelled to Sabah in North Borneo to spend [WO weeks searching for the wildlife of the region. Our group stayed in six locations, four of which yielded many species of phasmid. The rainy season had started early and therefore it frequently rained all afternoon, and often into the night.
    [Show full text]
  • CHAPTER 4: EPHEMEROPTERA (Mayflies)
    Guide to Aquatic Invertebrate Families of Mongolia | 2009 CHAPTER 4 EPHEMEROPTERA (Mayflies) EPHEMEROPTERA Draft June 17, 2009 Chapter 4 | EPHEMEROPTERA 45 Guide to Aquatic Invertebrate Families of Mongolia | 2009 ORDER EPHEMEROPTERA Mayflies 4 Mayfly larvae are found in a variety of locations including lakes, wetlands, streams, and rivers, but they are most common and diverse in lotic habitats. They are common and abundant in stream riffles and pools, at lake margins and in some cases lake bottoms. All mayfly larvae are aquatic with terrestrial adults. In most mayfly species the adult only lives for 1-2 days. Consequently, the majority of a mayfly’s life is spent in the water as a larva. The adult lifespan is so short there is no need for the insect to feed and therefore the adult does not possess functional mouthparts. Mayflies are often an indicator of good water quality because most mayflies are relatively intolerant of pollution. Mayflies are also an important food source for fish. Ephemeroptera Morphology Most mayflies have three caudal filaments (tails) (Figure 4.1) although in some taxa the terminal filament (middle tail) is greatly reduced and there appear to be only two caudal filaments (only one genus actually lacks the terminal filament). Mayflies have gills on the dorsal surface of the abdomen (Figure 4.1), but the number and shape of these gills vary widely between taxa. All mayflies possess only one tarsal claw at the end of each leg (Figure 4.1). Characters such as gill shape, gill position, and tarsal claw shape are used to separate different mayfly families.
    [Show full text]
  • Hickman's Pygmy Mountain Shrimp (Allanaspides Hickmani) Is a Small, Shrimp-Like Crustacean Belonging to the Family Anaspididae
    THREATENED SPECIES LISTING STATEMENT Hickman’s Pygmy Mountain Shrimp Allanaspides hickmani Swain Wilson and Ong 1971 Status Commonwealth Endangered Species Protection Act 1992 . .Not listed Tasmanian Threatened Species Protection Act 1995 . .Rare Description Hickman's pygmy mountain shrimp (Allanaspides hickmani) is a small, shrimp-like crustacean belonging to the family Anaspididae. This family contains three genera, Allanaspides, Anaspides and Paranaspides, all of which are restricted to Tasmania. Allanaspides species can be readily identified by the presence of a conspicuous transparent window on its back (dorsal window) and stalked eyes. The two species of Allanaspides (A. hickmani and A. helonomus) can be separated by the size, shape and colour of this window. Hickman's pygmy 6 mm mountain shrimp has a rectangular shaped dorsal window which covers most of the width of the dorsal surface Illustration: Karen Richards behind the head; the tissue below the dorsal window contains a bright red pigment. A. helonomus has a clear, oval-shaped dorsal window covering approximately half the width of the dorsal surface behind the head (Swain et al. 1970; Swain et al. 1971). Hickman's pygmy mountain shrimp has its eyes situated terminally on eyestalks and adult males attain a body length of 11.7 mm. A. helonomus has its eyes positioned laterally on the end of the eyestalks and is the slightly larger species, attaining a body length of 15 mm. A more detailed description of Hickman's pygmy mountain shrimp is provided by Swain et al. (1971). The life history of Hickman's pygmy mountain shrimp is poorly known and can only be inferred from what little information is available for A.
    [Show full text]
  • About the Book the Format Acknowledgments
    About the Book For more than ten years I have been working on a book on bryophyte ecology and was joined by Heinjo During, who has been very helpful in critiquing multiple versions of the chapters. But as the book progressed, the field of bryophyte ecology progressed faster. No chapter ever seemed to stay finished, hence the decision to publish online. Furthermore, rather than being a textbook, it is evolving into an encyclopedia that would be at least three volumes. Having reached the age when I could retire whenever I wanted to, I no longer needed be so concerned with the publish or perish paradigm. In keeping with the sharing nature of bryologists, and the need to educate the non-bryologists about the nature and role of bryophytes in the ecosystem, it seemed my personal goals could best be accomplished by publishing online. This has several advantages for me. I can choose the format I want, I can include lots of color images, and I can post chapters or parts of chapters as I complete them and update later if I find it important. Throughout the book I have posed questions. I have even attempt to offer hypotheses for many of these. It is my hope that these questions and hypotheses will inspire students of all ages to attempt to answer these. Some are simple and could even be done by elementary school children. Others are suitable for undergraduate projects. And some will take lifelong work or a large team of researchers around the world. Have fun with them! The Format The decision to publish Bryophyte Ecology as an ebook occurred after I had a publisher, and I am sure I have not thought of all the complexities of publishing as I complete things, rather than in the order of the planned organization.
    [Show full text]
  • Does Biogeography Have a Future in a Globalized World with Globalized Faunas?
    Contributions to Zoology, 77 (2) 127-133 (2008) Does biogeography have a future in a globalized world with globalized faunas? Frederick R. Schram Burke Museum, University of Washington, Seattle, P.O. Box 1567, Langley, WA 98260, USA, [email protected] ton.edu Key words: Anaspidacea, Bathynellacea, globalization, historical biogeography, vicariance Abstract in the Great Lakes and it is said that a new invader species is identified every eight months. The toll on The study of biogeography was once a pillar of evolution the fisheries of the Great Lakes alone has been dev- science. Both Darwin and especially Wallace found great in- astating. Another example is San Francisco Bay, spiration from the consideration of animal distributions. where some 234 invasive species have been recorded However, what is to happen to this discipline in a time of global trade, mass movement of people and goods, and the up to the present day, i.e., something like 90% of the resulting globalization of the planet’s biota? Can we still aquatic population of the bay. Finally, the infamous hope to delve into the fine points of past geography as it af- Chinese mitten crab, Eriocheir sinensis, is conduct- fected animal and plant evolution? Maybe we can, but only ing an on-going assault on Chesapeake Bay, and the with careful study of life forms that suffer minimal affects effect on the native blue crab populations is already – at present – from globalization, viz., marginal faunas of being measured. In the United States, invading ar- quite inaccessible environments. Two examples taken from syncarid crustaceans illustrate this point.
    [Show full text]