DOCSLIB.ORG

Agonist Lineages and Recognizes a Yeast-Derived TLR15 Is Unique To

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
Agonist Lineages and Recognizes a Yeast-Derived TLR15 Is Unique To TLR15 Is Unique to Avian and Reptilian Lineages and Recognizes a Yeast-Derived Agonist This information is current as Amy C. Boyd, Marylene Y. Peroval, John A. Hammond, of September 27, 2021. Michael D. Prickett, John R. Young and Adrian L. Smith J Immunol published online 12 October 2012 http://www.jimmunol.org/content/early/2012/10/12/jimmun ol.1101790 Downloaded from Why The JI? Submit online. • Rapid Reviews! 30 days* from submission to initial decision http://www.jimmunol.org/ • No Triage! Every submission reviewed by practicing scientists • Fast Publication! 4 weeks from acceptance to publication *average Subscription Information about subscribing to The Journal of Immunology is online at: by guest on September 27, 2021 http://jimmunol.org/subscription Permissions Submit copyright permission requests at: http://www.aai.org/About/Publications/JI/copyright.html Email Alerts Receive free email-alerts when new articles cite this article. Sign up at: http://jimmunol.org/alerts The Journal of Immunology is published twice each month by The American Association of Immunologists, Inc., 1451 Rockville Pike, Suite 650, Rockville, MD 20852 Copyright © 2012 by The American Association of Immunologists, Inc. All rights reserved. Print ISSN: 0022-1767 Online ISSN: 1550-6606. Published October 12, 2012, doi:10.4049/jimmunol.1101790 The Journal of Immunology TLR15 Is Unique to Avian and Reptilian Lineages and Recognizes a Yeast-Derived Agonist Amy C. Boyd,* Marylene Y. Peroval,† John A. Hammond,† Michael D. Prickett,‡ John R. Young,† and Adrian L. Smithx The TLRs represent a family of pattern recognition receptors critical in the induction of vertebrate immune responses. Between 10 and 13 different TLR genes can be identified in each vertebrate species, with many represented as orthologous genes in different species. The agonist specificity of orthologous TLR is also highly conserved. In contrast, TLR15 can only be identified in avian and reptilian genomes, suggesting that this receptor arose ∼320 million years ago after divergence of the bird/reptile and mammalian lineages. Transfection of a constitutively active form of chicken TLR15 led to NF-kB activation in HEK293 cells and induced cytokine mRNA upregulation in chicken cell lines. Full-length TLR15 mediated NF-kB induction in response to lysates from yeast, but not those derived from viral or bacterial pathogens, or a panel of well-characterized TLR agonists. TLR15 responses were Downloaded from induced by whole-cell lysates derived from Candida albicans, Saccharomyces cerevisiae, and Schizosaccharomyces pombe, but not zymosan preparations from S. cerevisiae. The ability of yeast lysate to activate TLR15-dependent NF-kB pathways (in transfection assays) or stimulate IL-1b mRNA upregulation in chicken macrophages was abrogated by heat inactivation or pre-exposure of the lysate to PMSF. Identification of yeast as an agonist source for TLR15 provides a functional framework for consideration of this TLR within the context of pattern recognition receptor evolution and may impact on the development of novel adjuvants. The Journal of Immunology, 2012, 189: 000–000. http://www.jimmunol.org/ attern recognition receptors (PRRs) are germline-encoded TLR families respond to distinct groups of agonists: the TLR1/2/6/ receptors that respond to a variety of conserved pathogen- 10 family recognizes lipopeptides (2, 3), the TLR3 family responds P associated molecular patterns (PAMPs) or microbial- to double-stranded RNA (4), the TLR4 family recognizes LPS (5), associated molecular patterns to initiate the inflammatory response. TLR5 family members respond to flagellin (6), and the TLR7–9 The TLRs are type I transmembrane glycoproteins characterized family recognizes nucleic acid motifs (7–10). by an agonist recognition domain comprising variable numbers of The sixth family, the TLR11 family (including TLR11–13 and leucine-rich repeats (LRRs) and the intracellular Toll/IL-1R ho- TLR20–23), is less well characterized and contains members that are mologous (TIR) domain, which mediates signaling. TLR genes and less widely distributed in vertebrates (1). Murine TLR11 (mTLR11) by guest on September 27, 2021 gene families are conserved in divergent species suggesting strong has been reported to respond to agonists derived from uropathogenic selective pressure for maintenance of function (1). The selective bacteria (11) and the apicomplexan protozoan PAMP, profilin (12). pressures to maintain TLR sequence are likely defined by the con- Pufferfish (Takifugu rubripes) TLR22 recognizes long double- servation of PAMPs (and damage-associated molecular patterns) and stranded RNA (13), and chicken TLR21 (cTLR21) has been re- the requirement to associate with accessory molecules (1). Between ported to respond to DNA rich in unmethylated CpG motifs (14, 15). 10 and 13 TLR genes are present in most vertebrates, and these can The diversity of agonist specificities in this family that includes be phylogenetically grouped into 6 major families (1). Five of the six nucleic acid and nonnucleic acid structures suggests that this family should be functionally subdivided. The cTLR repertoire comprises 10 genes including members of *The Jenner Institute, Nuffield Department of Clinical Medicine, University of Ox- each of the six families. Many cTLRs have been characterized ford, Oxford OX3 7DQ, United Kingdom; †Institute for Animal Health, Compton, Berkshire RG20 7NN, United Kingdom; ‡Cluster in Biomedicine, 34149 Basovizza, including TLR3, TLR4, TLR5, and TLR7, which are orthologous Trieste, Italy; and xDepartment of Zoology, University of Oxford, Oxford OX1 3PS, to their mammalian counterparts and share agonist specificity United Kingdom (16–21). Unlike mammals, chickens possess two TLR1 genes Received for publication July 5, 2011. Accepted for publication September 7, 2012. (TLR1.1 and 1.2) and two TLR2 genes (TLR2.1 and TLR2.2) that This work was supported by student support from the Biotechnology and Biological can form homodimers or heterodimers and respond to peptido- Sciences Research Council and the Institute for Animal Health, with additional support from Department for Environment, Food and Rural Affairs Project OD0547. A.L.S. is glycan, diacylated lipopeptides, and triacylated lipopeptides (22– a fellow of The Jenner Institute. 24). In mammalian species, these agonists are recognized by Address correspondence and reprint requests to Dr. Adrian L. Smith, Department of various heterodimers composed of TLR1 or TLR6 with the single Zoology, University of Oxford, Tinbergen Building, South Parks Road, Oxford OX1 TLR2. The duplication of TLR2 in the chicken and zebra finch 3PS, U.K. E-mail address: [email protected] (22, 25) appears to have occurred before the divergence of birds Abbreviations used in this article: ALV, avian leukosis virus; ATCC, American Type and mammals, as TLR2 pseudogenes can be found upstream of Culture Collection; CA, constitutively active; CAcTIR4, constitutively active chicken TLR4; CAcTIR15, constitutively active chicken TLR15; CAmTIR4, constitutively the functional TLR2 sequences in opossum, dog, and human (1). active murine TLR4; CATLR, constitutively active TLR; Ct, threshold cycle; cTLR, Additional differences between mammalian and avian TLR rep- chicken TLR; GOI, gene of interest; LRR, leucine-rich repeat; mTIRcon, murine TIR deficient control construct; mTLR, murine TLR; mya, million years ago; PAMP, ertoires include the disruption of TLR8 in galliform and anseri- pathogen-associated molecular pattern; PRR, pattern recognition receptor; qRT-PCR, form birds by a retroviral insertion (17, 26) and the lack of a TLR9 quantitative RT-PCR; SEAP, secreted human embryonic alkaline phosphatase; TIR, homolog in the chicken. Before this study, TLR15 was identified Toll/IL-1R homologous. in the chicken and zebra finch genomes (1, 25). We identified Copyright Ó 2012 by The American Association of Immunologists, Inc. 0022-1767/12/$16.00 TLR15 in the genomic assemblies of two further avian species, www.jimmunol.org/cgi/doi/10.4049/jimmunol.1101790 2 EVOLUTION AND FUNCTION OF TLR15 the duck (Anas platyrhynchos) and wild turkey (Meleagris gal- CCACCCAATCCAGGAAATGTTAACCCA-39. Primers were synthesized lopavo). A partial TLR15 sequence was also identified in the by Sigma. Probes labeled with the fluorescent reporter dye FAM at the 59 9 genome of the Carolina anole lizard (Anolis carolinensis). These end and the quencher TAMRA at the 3 end were obtained from Euro- gentec (Southampton, U.K.). Primer and probe sequences for 28S RNA sequences afforded an opportunity to resolve the phylogenetic and cytokine mRNA assays were as previously published (32, 33). relationship between TLR15 and the other TLR families. Fur- Quantitative RT-PCR (qRT-PCR) was performed using the Applied thermore, we demonstrated the functional capability of TLR15: Biosystems TaqMan FAST Universal PCR Master Mix. Amplification and it activates innate immunity and specifically recognizes novel detection of target sequences were performed using the Applied Biosystems 7500 FAST Real-Time PCR System with the following cycle conditions: agonist structures present in whole-cell lysates derived from one cycle of 48˚C for 30 min and 94˚C for 20 s, and 40 cycles of 94˚C for yeast/fungal pathogens. 3 s, 60˚C for 30 s. For analysis of cTLR mRNA expression in tissues, 28S values were used Materials and Methods to adjust for differences in input RNA. Samples were diluted 1:5 for gene of interest (GOI) and 1:500 for 28S reactions. For analysis of cTLR mRNA
Load more

Recommended publications
  • Reptiles A. Cladistics 1. Many Groups of Organisms
    Reptiles A. Cladistics 1. Many groups of organisms are “polyphyletic” a. This means that the group combines 2 or more lineages - example=fish 2. Cladistics follows only pure lineages going back in time - example Osteichthys B. Reptile Classifiecation - looks like a polyphyletic group 1. Dry skin - no loss of water through skin like amphibians 2. Aminotic egg - an egg that can survive on dry land - in contrast with the amphibian egg C. Mammals and Birds are derived from different lineages of reptiles (We will see below) D. Stem Reptiles 1. Different lineages based on the temporal region of their skulls - number of holes (or bars) a. These holes are necessary to accommodate large jaw muscles b. Anapsid Skull - no holes in temporal - jaws can move fast, but with little force 1. Muscles that move the jaw are small 2. There is no good paleotological evidence for the transition between amphibians and reptiles - no fossil intermediates a. Fossil amphibians have lots of dermal bones in skull b. Amphibians have no temporal openings in skull 1. (Aside) both fossil amphibians and primitive reptiles have a parietal “eye” that senses light and dark (“third” eye in middle of head) c. Reptile skull is higher than amphibian to accomodate larger jaw muscles d. Of the modern reptiles only turtles are anapsids 2. Diapsid Skull - has holes in the temporal region a. Diapsid reptiles gave rise to lizards and snakes - they have a diapsid skull 1. Also Tuatara, crocodiles, dinosaurs and pterydactyls Reptiles b. One group of diapsids also had a pre-orbital hole in the skull in front of eye - this hole is still preserved in the birds - this anatomy suggests strongly that the birds are derived from the diapsid reptiles 3.
    [Show full text]
  • Morphology, Phylogeny, and Evolution of Diadectidae (Cotylosauria: Diadectomorpha)
    Morphology, Phylogeny, and Evolution of Diadectidae (Cotylosauria: Diadectomorpha) by Richard Kissel A thesis submitted in conformity with the requirements for the degree of doctor of philosophy Graduate Department of Ecology & Evolutionary Biology University of Toronto © Copyright by Richard Kissel 2010 Morphology, Phylogeny, and Evolution of Diadectidae (Cotylosauria: Diadectomorpha) Richard Kissel Doctor of Philosophy Graduate Department of Ecology & Evolutionary Biology University of Toronto 2010 Abstract Based on dental, cranial, and postcranial anatomy, members of the Permo-Carboniferous clade Diadectidae are generally regarded as the earliest tetrapods capable of processing high-fiber plant material; presented here is a review of diadectid morphology, phylogeny, taxonomy, and paleozoogeography. Phylogenetic analyses support the monophyly of Diadectidae within Diadectomorpha, the sister-group to Amniota, with Limnoscelis as the sister-taxon to Tseajaia + Diadectidae. Analysis of diadectid interrelationships of all known taxa for which adequate specimens and information are known—the first of its kind conducted—positions Ambedus pusillus as the sister-taxon to all other forms, with Diadectes sanmiguelensis, Orobates pabsti, Desmatodon hesperis, Diadectes absitus, and (Diadectes sideropelicus + Diadectes tenuitectes + Diasparactus zenos) representing progressively more derived taxa in a series of nested clades. In light of these results, it is recommended herein that the species Diadectes sanmiguelensis be referred to the new genus
    [Show full text]
  • The Global Distribution of Tetrapods Reveals a Need for Targeted Reptile
    1 The global distribution of tetrapods reveals a need for targeted reptile 2 conservation 3 4 Uri Roll#1,2, Anat Feldman#3, Maria Novosolov#3, Allen Allison4, Aaron M. Bauer5, Rodolphe 5 Bernard6, Monika Böhm7, Fernando Castro-Herrera8, Laurent Chirio9, Ben Collen10, Guarino R. 6 Colli11, Lital Dabool12 Indraneil Das13, Tiffany M. Doan14, Lee L. Grismer15, Marinus 7 Hoogmoed16, Yuval Itescu3, Fred Kraus17, Matthew LeBreton18, Amir Lewin3, Marcio Martins19, 8 Erez Maza3, Danny Meirte20, Zoltán T. Nagy21, Cristiano de C. Nogueira19, Olivier S.G. 9 Pauwels22, Daniel Pincheira-Donoso23, Gary Powney24, Roberto Sindaco25, Oliver Tallowin3, 10 Omar Torres-Carvajal26, Jean-François Trape27, Enav Vidan3, Peter Uetz28, Philipp Wagner5,29, 11 Yuezhao Wang30, C David L Orme6, Richard Grenyer✝1 and Shai Meiri✝*3 12 13 # Contributed equally to the paper 14 ✝ Contributed equally to the paper 15 * Corresponding author 16 17 Affiliations: 18 1 School of Geography and the Environment, University of Oxford, Oxford, OX13QY, UK. 19 2 Mitrani Department of Desert Ecology, The Jacob Blaustein Institutes for Desert Research, 20 Ben-Gurion University, Midreshet Ben-Gurion 8499000, Israel. (Current address) 21 3 Department of Zoology, Tel-Aviv University, Tel-Aviv 6997801, Israel. 22 4 Hawaii Biological Survey, 4 Bishop Museum, Honolulu, HI 96817, USA. 23 5 Department of Biology, Villanova University, Villanova, PA 19085, USA. 24 6 Department of Life Sciences, Imperial College London, Silwood Park Campus Silwood Park, 25 Ascot, Berkshire, SL5 7PY, UK 26 7 Institute of Zoology, Zoological Society of London, London NW1 4RY, UK. 27 8 School of Basic Sciences, Physiology Sciences Department, Universidad del Valle, Colombia.
    [Show full text]
  • Marine Reptiles Arne R
    Virginia Commonwealth University VCU Scholars Compass Study of Biological Complexity Publications Center for the Study of Biological Complexity 2011 Marine Reptiles Arne R. Rasmessen The Royal Danish Academy of Fine Arts John D. Murphy Field Museum of Natural History Medy Ompi Sam Ratulangi University J. Whitfield iG bbons University of Georgia Peter Uetz Virginia Commonwealth University, [email protected] Follow this and additional works at: http://scholarscompass.vcu.edu/csbc_pubs Part of the Life Sciences Commons Copyright: © 2011 Rasmussen et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Downloaded from http://scholarscompass.vcu.edu/csbc_pubs/20 This Article is brought to you for free and open access by the Center for the Study of Biological Complexity at VCU Scholars Compass. It has been accepted for inclusion in Study of Biological Complexity Publications by an authorized administrator of VCU Scholars Compass. For more information, please contact [email protected]. Review Marine Reptiles Arne Redsted Rasmussen1, John C. Murphy2, Medy Ompi3, J. Whitfield Gibbons4, Peter Uetz5* 1 School of Conservation, The Royal Danish Academy of Fine Arts, Copenhagen, Denmark, 2 Division of Amphibians and Reptiles, Field Museum of Natural History, Chicago, Illinois, United States of America, 3 Marine Biology Laboratory, Faculty of Fisheries and Marine Sciences, Sam Ratulangi University, Manado, North Sulawesi, Indonesia, 4 Savannah River Ecology Lab, University of Georgia, Aiken, South Carolina, United States of America, 5 Center for the Study of Biological Complexity, Virginia Commonwealth University, Richmond, Virginia, United States of America Of the more than 12,000 species and subspecies of extant Caribbean, although some species occasionally travel as far north reptiles, about 100 have re-entered the ocean.
    [Show full text]
  • Meet the Herps!
    Science Standards Correlation SC06-S2C2-03, SC04-S4C1-04, SC05-S4C1-01, SC04-S4C1-06, SC07-S4C3-02, SC08- S4C4-01, 02&06 MEET THE HERPS! Some can go without a meal for more than a year. Others can live for a century, but not really reach a ripe old age for another couple of decades. One species is able to squirt blood from its eyes. What kinds of animals are these? They’re herps – the collective name given to reptiles and amphibians. What Is Herpetology? The word “herp” comes from the word “herpeton,” the Greek word for “crawling things.” Herpetology is the branch of science focusing on reptiles and amphibians. The reptiles are divided into four major groups: lizards, snakes, turtles, and crocodilians. Three major groups – frogs (including toads), salamanders and caecilians – make up the amphibians. A herpetologist studies animals from all seven of these groups. Even though reptiles and amphibians are grouped together for study, they are two very different kinds of animals. They are related in the sense that early reptiles evolved from amphibians – just as birds, and later mammals, evolved from reptiles. But reptiles and amphibians are each in a scientific class of their own, just as mammals are in their own separate class. One of the reasons reptiles and amphibians are lumped together under the heading of “herps” is that, at one time, naturalists thought the two kinds of animals were much more closely related than they really are, and the practice of studying them together just persisted through the years. Reptiles vs. Amphibians: How Are They Different? Many of the differences between reptiles and amphibians are internal (inside the body).
    [Show full text]
  • A New Species of the Sauropsid Reptile Nothosaurus from the Lower Muschelkalk of the Western Germanic Basin, Winterswijk, the Netherlands
    A new species of the sauropsid reptile Nothosaurus from the Lower Muschelkalk of the western Germanic Basin, Winterswijk, The Netherlands Nicole Klein and Paul C.H. Albers Acta Palaeontologica Polonica 54 (4), 2009: 589-598 doi: http://dx.doi.org/10.4202/app.2008.0083 A nothosaur skull recently discovered from the Lower Muschelkalk (early Anisian) locality of Winterswijk, The Netherlands, represents at only 46 mm in length the smallest nothosaur skull known today. It resembles largely the skull morphology of Nothosaurus marchicus. Differences concern beside the size, the straight rectangular and relative broad parietals, the short posterior extent of the maxilla, the skull proportions, and the overall low number of maxillary teeth. In spite of its small size, the skull can not unequivocally be interpreted as juvenile. It shows fused premaxillae, nasals, frontals, and parietals, a nearly co−ossified jugal, and fully developed braincase elements, such as a basisphenoid and massive epipterygoids. Adding the specimen to an existing phylogenetic analysis shows that it should be assigned to a new species, Nothosaurus winkelhorsti sp. nov., at least until its juvenile status can be unequivocally verified. Nothosaurus winkelhorsti sp. nov. represents, together with Nothosaurus juvenilis, the most basal nothosaur, so far. Key words: Sauropterygia, Nothosaurus, ontogeny, Anisian, The Netherlands. Nicole Klein [[email protected]], 1Steinmann Institute, Paleontology, University of Bonn, Nußallee 8, 53115 Bonn, Germany; Paul C.H. Albers [[email protected]], Naturalis, Leiden, The Netherlands. Darwinweg 2, 2333 CR Leiden, The Netherlands; This is an open-access article distributed under the terms of the Creative Commons Attribution License (for details please see creativecommons.org), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
    [Show full text]
  • 190 World's First Herbivorous Filter-Feeding Marine Reptile
    BCAS Vol.30 No.3 2016 Earth Sciences World’s First Herbivorous Filter- feeding Marine Reptile ome strange creatures cropped up in the wake Its head was poorly preserved, but it seemed to have of one of Earth’s biggest mass extinctions a flamingo-like beak. However, in a paper published S252 million years ago. In 2014, scientists May 6 in Science Advances, Dr. LI Chun, Institute of discovered a bizarre fossil – a crocodile-sized sea- Vertebrate Paleontology and Paleoanthropology (IVPP), dwelling reptile, Atopodentatus unicus, that lived 242 Chinese Academy of Sciences, and his international million years ago in what today is southwestern China. team described two new specimens and revealed what Fossil and reconstruction of Atopodentatus unicus (Image by IVPP) 190 Bulletin of the Chinese Academy of Sciences Vol.30 No.3 2016 Science Watch Earth Sciences was really going on—that "beak" is actually part of a among marine reptiles. It is older than other marine hammerhead-shaped jaw apparatus, which the reptile used animals that ate plants with a filter-feeding system by to feed on plants on the ocean floor. It's the earliest known about eight million years, said the team. example of an herbivorous marine reptile. Atopodentatus appeared during the Triassic period These two newly discovered specimens of soon after the biggest mass extinction of species in Earth's Atopodentatus were collected from the Middle Triassic history, illustrating that life recovered and diversified (Anisian) Guanling Formation of Luoping County, Yunnan more quickly than previously thought. Other oddball Province, southwestern China. The new specimens clearly creatures also swam the seas at the time, including a demonstrate that rather than being downturned, the reptile called Dinocephalosaurus whose neck comprised rostrum developed into a “hammerhead” with pronounced half of its 17-foot (5.25 meters) length.
    [Show full text]
  • The Vertebrate Fauna of the New Mexico Permian Alfred S
    New Mexico Geological Society Downloaded from: http://nmgs.nmt.edu/publications/guidebooks/11 The vertebrate fauna of the New Mexico Permian Alfred S. Romer, 1960, pp. 48-54 in: Rio Chama Country, Beaumont, E. C.; Read, C. B.; [eds.], New Mexico Geological Society 11th Annual Fall Field Conference Guidebook, 129 p. This is one of many related papers that were included in the 1960 NMGS Fall Field Conference Guidebook. Annual NMGS Fall Field Conference Guidebooks Every fall since 1950, the New Mexico Geological Society (NMGS) has held an annual Fall Field Conference that explores some region of New Mexico (or surrounding states). Always well attended, these conferences provide a guidebook to participants. Besides detailed road logs, the guidebooks contain many well written, edited, and peer-reviewed geoscience papers. These books have set the national standard for geologic guidebooks and are an essential geologic reference for anyone working in or around New Mexico. Free Downloads NMGS has decided to make peer-reviewed papers from our Fall Field Conference guidebooks available for free download. Non-members will have access to guidebook papers two years after publication. Members have access to all papers. This is in keeping with our mission of promoting interest, research, and cooperation regarding geology in New Mexico. However, guidebook sales represent a significant proportion of our operating budget. Therefore, only research papers are available for download. Road logs, mini-papers, maps, stratigraphic charts, and other selected content are available only in the printed guidebooks. Copyright Information Publications of the New Mexico Geological Society, printed and electronic, are protected by the copyright laws of the United States.
    [Show full text]
  • Snake and Lizards of Minnesota
    SNAKES AND LIZARDS OF MINNESOTA TABLE OF CONTENTS Acknowledgments . 4 Introduction . 6 Key to Minnesota’s Snakes . 24 Common Gartersnake . 26 Common Watersnake . 28 DeKay’s Brownsnake . 30 Eastern Hog‑nosed Snake . 32 Gophersnake . 34 Lined Snake . 36 Massasauga . 38 Milksnake . 40 North American Racer . 42 Plains Gartersnake . 44 Plains Hog‑nosed Snake . 46 Red‑bellied Snake . 48 Ring‑necked Snake . 50 Smooth Greensnake . 52 Timber Rattlesnake . 54 Western Foxsnake . 56 Western Ratsnake . 58 Key to Minnesota’s Lizards . 61 Common Five‑lined Skink . .. 62 Prairie Skink . 64 Six‑lined Racerunner . 66 Glossary . 68 Appendix . 70 Help Minnesota’s Wildlife! . 71 Cover photos: Timber rattlesnakes photograph by Barb Perry . Common five‑lined skink photograph by Carol Hall . Left: Park naturalist holding gophersnake . Photograph by Deborah Rose . ACKNOWLEDGMENTS Text Rebecca Christoffel, PhD, Contractor Jaime Edwards, Department of Natural Resources (DNR) Nongame Wildlife Specialist Barb Perry, DNR Nongame Wildlife Technician Snakes and Lizards Design of Minnesota Creative Services Unit, DNR Operation Services Division Editing Carol Hall, DNR Minnesota Biological Survey (MBS), Herpetologist Liz Harper, DNR Ecological and Water Resources (EWR), Assistant Central Regional Manager Erica Hoagland, DNR EWR, Nongame Wildlife Specialist Tim Koppelman, DNR Fish and Wildlife, Assistant Area Wildlife Manager Jeff LeClere, DNR, MBS, Animal Survey Specialist John Moriarity, Senior Manager of Wildlife, Three Rivers Park District Pam Perry, DNR, EWR, Nongame Wildlife Lake Specialist (Retired) This booklet was funded through a State Wildlife Grant and the Nongame Wildlife Program, DNR Ecological and Water Resources Division . Thank you for your contributions! See inside back cover . ECOLOGICAL AND WATER RESOURCES INTRODUCTION is understandable in Minnesota, spend most of the active season .
    [Show full text]
  • Amniota: Eureptilia) from the Upper Permian of Mallorca (Balearic Islands, Western Mediterranean)
    A large multiple tooth-rowed captorhinid reptile (Amniota: Eureptilia) from the Upper Permian of Mallorca (Balearic Islands, Western Mediterranean) TORSTEN LIEBRECHT,1 JOSEP FORTUNY,2 ÀNGEL GALOBART,2 JOHANNES MÜLLER,1 and P. MARTIN SANDER,3 1 Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Invalidenstraße 43, 10115 Berlin, Germany, [email protected], [email protected]; 2 Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, Carrer de les Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona, Spain, [email protected], [email protected]; 3 Steinmann-Institut für Geologie, Mineralogie und Paläontologie, Rheinische Friedrich-Wilhelms-Universität Bonn, Nussallee 8, 53115 Bonn, Germany, [email protected] Supplementary data GEOLOGIC SETTING Today, the Balearic Islands (Spanish: Islas Baleares; Catalan: Illes Balears; Fig. S1) represent the geomorphologically highest, and thus emergent, parts of the north-eastern extension of the Betic Cordillera of southern Spain. This extension is also called the Balearic Promontory, which to the northwest is separated from the Iberian mainland by the Valencia Trough, an Oligocene to recent extensional structure that has a complex tectonic history closely connected to the Alpidic collisional movements that affected the western Mediterranean realm in the Late Mesozoic and Cenozoic (Roca, 1996). The northwestern part of the Island of Mallorca is occupied by the Serra de Tramuntana (Serra del Nord), a southwest-northeast trending horst-like structure that internally is built up of southeast-dipping Alpidic thrust sheets of unmetamorphosed, predominantly calcareous rocks of Jurassic age. Exposures of Permian to Middle Triassic terrestrial redbeds (so-called ‘Permo- Triassic’) are present only at the northwestern flank of the Serra de Tramuntana in the coastal area between the villages of Estellencs and Valldemosa.
    [Show full text]
  • Reptile Observation Observe Reptile Characteristics and Behaviors at the Zoo
    Reptile Observation Observe reptile characteristics and behaviors at the zoo Grade 2 Objectives • Students will learn about reptiles through observation and drawing conclusions. Materials • provided zoo map Background Information • All reptiles have a backbone, breathe with lungs, and lay dry, scaly skin. • Reptiles lay eggs that have a dry leathery shell. Unlike amphibians, which have eggs with a jelly coat Key Words that must remain in water, reptiles are less dependent on water. • reptile • Basic Reptile Types • habitiat - Turtles are the only reptiles that have their “houses” on their backs. • environment - Lizards have visible, moving eyelids, limbs, and an ear opening on each side of their head. - Snakes lack limbs, eyelids, and ear openings. - Crocodilians include alligators as well as crocodiles. They are the most ancestral standards of the reptiles. • SCI.2.3.1 • SCI.2.3.2 Recommended Assessment • Have a group discussion about the observations made by students. Extensions • Assign each child or group a Reptile Treasure to discover as they move through zoo; for example, the heaviest reptile, the smallest reptile, the longest reptile, a reptile with three colors, etc. The children should report on the results of their treasure hunt when they get back to school. Reptile Observation Observe the following reptiles in Dr. Diversity’s Rain Forest Research Station while you are at the zoo. Fill in your observation on the chart. Name Timor Name of Komodo Reticulated Monitor Reptile Dragon Python Lizard Kind of snake snake snake Reptile lizard lizard lizard turtle turtle turtle crocodilian crocodilian crocodilian What it Looks Like What it Eats Interesting Facts Reptile Observation Observe the following reptiles in Dr.
    [Show full text]
  • Amniotes a Closer Look O Si a L O N Nalysis Es Gy G 7A, 7B, 10A G Athers
    DO NOT EDIT--Changes must be made through “File info” CorrectionKey=B A Closer Look CHAPTER 26 at Amniotes BIG IDEA Reptiles, birds, and mammals all share similar traits, but other, specialized characteristics classify these animals into different groups. 26.1 Amniotes 7B, 10A Data Analysis CHOOSING GRAPHS 2G 26.2 Reptiles 7A, 7B, 8C, 10A 26.3 Birds 7A, 7B, 10A 26.4 Mammals 7A, 7B, 7D, 8B, 8C,10A ONLINE BIOLOGY HMDScience.com ONLINE Labs ■■ The Effect of Temperature on Cold-Blooded ■■ QuickLab Comparing Feathers Organisms ■■ A Bird’s Airframe ■■ Video Lab Bird Digestion ■■ The Parts of an Egg ■■ Video Lab Mammalian Characteristics ■■ Migration and Range ■■ Form and Function of Teeth ■■ Identifying Features: Hair (t) ©F. Stuart Westmorland/Photo Researchers, Inc. Researchers, Westmorland/Photo Stuart (t) ©F. 754 Unit 8: Animals DO NOT EDIT--Changes must be made through “File info” CorrectionKey=B Q Is this a monkey or a mouse? The large eyes and ears of this eastern tarsier make it an excellent nocturnal hunter. These unusual mammals are primates and are closely related to modern monkeys. About the same size as a kitten, a tarsier has strong hind legs similar to those of frogs. The tarsier’s eyes cannot move but it has a full range of view because it can rotate its head almost 360 degrees. READING TOOLBOX This reading tool can help you learn the material in the following pages. USING LANGUAGE YOUR TURN Word Problems A good way to begin solving a word Refer to the word problem example to answer the problem is to figure out what it is asking for.
    [Show full text]