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 Monitoring in
Neurocritical Care
 Monitoring in
Neurocritical Care

Peter D. le Roux, MD, FACS

Associate Professor of Neurosurgery
Perelman School of Medicine at the University of
Pennsylvania;
Department of Neurosurgery
Pennsylvania Hospital
Philadelphia, Pennsylvania

Joshua M. Levine, MD
Assistant Professor
Departments of Neurology, Neurosurgery, and
Anesthesiology and Critical Care
Perelman School of Medicine at the University of
Pennsylvania;
Co-Director, Neurointensive Care Unit
Hospital of the University of Pennsylvania
Philadelphia, Pennsylvania

W. Andrew Kofke, MD, MBA, FCCM

Professor
Departments of Anesthesiology and Critical Care and
Neurosurgery
Perelman School of Medicine at the University of
Pennsylvania;
Director, Neuroanesthesia
Department of Anesthesiology and Critical Care
University of Pennsylvania Health System;
Co-Director, Neurointensive Care Unit
Hospital of the University of Pennsylvania
Philadelphia, Pennsylvania
1600 John F. Kennedy Blvd.
Ste 1800
Philadelphia, PA 19103-2899

MONITORING IN NEUROCRITICAL CARE ISBN: 978-1-4377-0167-8

Copyright © 2013 by Saunders, an imprint of Elsevier Inc.
Chapter 22: “Renal, Electrolyte, and Acid Base Assessment”: Guy M. Dugan retains copyright to his original
contribution.
Chapter 23: “Gastrointestinal and Hepatic Disorders” by Christiana E. Hall and Aashish R. Patel:
Christiana E. Hall retains copyright to her original portion of the contribution only.

No part of this publication may be reproduced or transmitted in any form or by any means, electronic or
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permission in writing from the Publisher. Details on how to seek permission, further information about the
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This book and the individual contributions contained in it are protected under copyright by the Publisher
(other than as may be noted herein).

Notices

Knowledge and best practice in this field are constantly changing. As new research and experience broaden
our understanding, changes in research methods, professional practices, or medical treatment may become
necessary.
Practitioners and researchers must always rely on their own experience and knowledge in evaluating
and using any information, methods, compounds, or experiments described herein. In using such
information or methods, they should be mindful of their own safety and the safety of others, including
parties for whom they have a professional responsibility.
With respect to any drug or pharmaceutical products identified, readers are advised to check the most
current information provided (i) on procedures featured or (ii) by the manufacturer of each product to be
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and contraindications. It is the responsibility of practitioners, relying on their own experience and
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To the fullest extent of the law, neither the Publisher nor the authors, contributors, or editors, assume
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contained in the material herein.

Library of Congress Cataloging-in-Publication Data

Monitoring in neurocritical care / [edited by] Peter D. Le Roux, Joshua M. Levine, W. Andrew Kofke.
    p. ; cm.
   Includes bibliographical references and index.
   ISBN 978-1-4377-0167-8 (hardcover : alk. paper)
   I. Le Roux, Peter D. II. Levine, Joshua M. III. Kofke, W. Andrew.
   [DNLM:1. Monitoring, Intraoperative–methods. 2. Neurosurgical Procedures. 3. Central Nervous
System Diseases–surgery. 4. Intensive Care–methods. 5. Intensive Care Units–organization &
administration. WL 368]
   616.02’8–dc23
2012036807

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To the ICU nurses who take care of all our patients
Preface
Neurocritical care has evolved rapidly in the past 10 years. understanding of what information the technology provides.
It is a specialty that focuses on the critical care manage- These advances have been chronicled in numerous contribu-
ment of patients with catastrophic neurologic diseases, tions to the scientific literature, the sheer volume of which
including primary neurologic pathologies such as traumatic makes it difficult for healthcare providers and device engi-
brain injury (TBI), ischemic stroke, intracerebral hemorrhage neers to keep up to date with the knowledge necessary to
(ICH), subarachnoid hemorrhage (SAH), brain tumors, provide the best patient care. In addition, although several
infection (e.g., HIV, TB, meningitis), spinal cord injury, and textbooks on critical care or head injury briefly discuss moni-
acute ascending neuropathies. In addition, neurologic dys- toring in a chapter or two, there is a paucity of information
function occurs in many diverse systemic disorders, including that summarizes all aspects of neuromonitoring and no text-
hypoxia (e.g., post cardiac arrest, near drowning), liver dys- book that is dedicated to monitoring in neurocritical care.
function, electrolyte abnormalities, high altitude sickness, This book, Monitoring in Neurocritical Care, represents a
eclampsia, lead intoxication, and malignant hypertension. comprehensive review of neuromonitoring. We have designed
There are many reasons why brain injury or damage occurs this textbook to provide the reader with a practical but in-
in patients with neurologic disorders. In particular, multiple depth reference that describes the scientific basis and rationale
experimental and clinical studies have demonstrated a close for use of a particular monitor, the information it provides,
relationship between variables such as hypoxia, increased and how this information can be used to manage the neuro-
intracranial pressure, arterial hypotension, hyperglycemia, critical care patient in an integrated fashion. We have been
fever, and seizures with neurologic outcome, and accumulat- fortunate to have chapters written by authors from around the
ing evidence suggests that brain damage evolves over time. world. The contributors are clinicians, engineers, information
Minimizing the burden of this delayed, or “secondary,” brain technology experts, and researchers who have extensive expe-
injury, has become the focus of modern neurocritical care. rience in the field, and each has provided an excellent and
However, despite much research, trials in neuroprotection timely review. We hope that the reader will gain a comprehen-
have largely failed, in part because of their association with sive understanding about neuromonitoring and neurocritical
prognostic heterogeneity, multiple mechanisms of cellular care and an insight into existent controversies and potential
damage, and a paucity of early mechanistic endpoints. This future management. Its contents will be relevant to neurolo-
has led to a realization that strategies of care, or “bundles,” gists, neurosurgeons, neuroanesthesiologists, neurointensiv-
rather than single agents, and approaches that are tailored to ists, and neuroscience nurses, and will serve as a useful resource
individual patient physiology and pathophysiology are neces- to intensivists working in medical and surgical ICUs. For those
sary to improve outcome. who are interested in clinical or laboratory research on brain
In daily practice neurointensivists focus on the recognition injury in its broad sense, this book will provide many ideas
of subtle changes in the neurologic condition, interactions and references and will be a stepping-stone to further progress
between the brain and systemic derangements, and brain in understanding a complex problem. The text also will serve
physiology. The challenge for intensivists today is to identify as a reference and guide for many engineers, bioengineers, and
individuals who are at risk of developing disease or secondary computer experts who work on medical device and bioinfor-
injury, determine disease severity, and distinguish responders matics development.
from nonresponders to therapy (i.e., individualized and tar- We have divided the book into seven sections. Section I,
geted medicine). Monitoring is one tool that may answer these Background, provides information about cerebral metabo-
challenges, and it has become central to the management of lism, the principles of neurocritical care, informatics, quality
secondary brain injury and to individualized care. In recent assessment, the role of ICU design and nursing, specific con-
years (interestingly in concert with the evolution of neuro- siderations in children, the effects of anesthetic agents on
critical care as a distinct specialty), technology developments monitors, and a discussion on the relationship between bio-
have resulted in several new monitoring techniques that ethics and monitoring. Section II, Clinical and Laboratory
provide the neurointensivist with information about brain Assessment, reviews clinical evaluation, sedation, pain, delir-
and cellular function. Techniques to better monitor function ium, outcomes such as neuropsychological and brain death,
of the heart, lung, liver, kidney, and blood also have evolved. extracerebral organ systems, laboratory analysis, and the role
In addition, when the various techniques are combined (“mul- of biomarkers. Section III, Electrophysiology, is devoted to
timodal monitoring”), a more accurate overall picture of brain evoked potentials and electroencephalography. Section IV,
function is produced. This approach, along with new com- Radiology, discusses the use and integration of various tech-
puter systems that integrate data at the bedside, and the niques, including computed tomography, xenon-CT, MRI,
emerging field of bioinformatics may change the way patients PET, and SPECT in neurocritical care. Section V, Cerebral
with brain injury are managed in the future. Blood Flow, is a review of techniques such as neurosonology,
In the last decade, there have been many advances in laser Doppler flowmetry, thermal diffusion flowmetry, jugular
neurocritical care monitoring technology and a better bulb oximetry, and near infrared spectroscopy. Section VI,
vii
viii Preface

Intracranial Monitoring, provides an in-depth review of We would like to express our appreciation and acknowledge
invasive techniques, including intracranial pressure, brain the efforts of all contributors to this volume. We also thank
oxygen, cerebral microdialysis, and brain temperature. The the editorial, design, and production staff at Elsevier Science,
final section, Computers, Engineering, and the Future, pro- in particular Janice Gaillard, Charlotta Kryhl, Julie Goolsby,
vides a description of device development, engineering, sim- Kate Crowley, Angela Rufino, Louis Forgione, Cheryl Abbott,
ulation, telemedicine, robotics, information processing, data Louise King, and Anne Altepeter, who have been very helpful
acquisition and storage, medical informatics and multimo- in producing this volume, and Yolanda Caban who provided
dality monitoring, noninvasive brain monitoring, and a dis- excellent administrative assistance. Finally, we thank Barbara
cussion of potential future developments. It is important for Williams who provided outstanding editorial assistance and
the reader to realize that the “ideal brain monitor” does not made this book possible.
yet exist and no single monitor will by itself affect outcome. Peter D. le Roux, MD, FACS
Instead, it is the information provided by a monitor and how Joshua M. Levine, MD
we as healthcare providers interpret and apply the informa- W. Andrew Kofke, MD, MBA, FCCM
tion that has the potential to improve outcome and lead to
new insights into disease processes.
Contributors
Pippa G. Al-Rawi, BSc Rosette C. Biester, PhD Maurizio Cereda, MD
Research Associate Polytrauma Neuropsychologist Assistant Professor
Neurosurgery Unit Department of Behavioral Health Department of Anesthesiology and Critical Care
Department of Clinical Neurosciences Philadelphia Veterans Affairs Medical Center; Perelman School of Medicine at the University
University of Cambridge/Addenbrooke’s Auxiliary Health Care Provider of Pennsylvania
Hospital Department of Physical Medicine and Philadelphia, Pennsylvania
Cambridge, United Kingdom Rehabilitation
University of Pennsylvania Health System Randall M. Chesnut, MD, FCCM, FACS
Pamela J. Amelung, MD Philadelphia, Pennsylvania Integra Endowed Professor of Neurotrauma
Clinical Associate Professor Department of Neurological Surgery
Department of Medicine Peter M. Black, MD, PhD Department of Orthopaedic Surgery
University of Maryland School of Medicine; Center for Advanced Brain and Spine Surgery Adjunct Professor, School of Global Health
Physician Liaison Natick, Massachusetts; Harborview Medical Center, University of
Philips VISICU Professor of Neurosurgery Washington
Baltimore, Maryland Department of Surgery Seattle, Washington
Harvard Medical School
Michal Arkuszewski, MD, PhD Boston, Massachusetts Jan Claassen, MD
Department of Neurology, Central University Attending Neurointensivist
Hospital Thomas P. Bleck, MD, FCCM Department of Neurology
Medical University of Silesia Professor Division of Neurocritical Care
Katowice, Poland; Departments of Neurological Sciences, Columbia University College of Physicians and
Department of Radiology, Neuroradiology Neurosurgery, Anesthesiology, and Medicine Surgeons
Division Rush Medical College; New York, New York
Perelman School of Medicine at the University Associate Chief Medical Officer for Critical Care
of Pennsylvania Rush University Medical Center Wendy A. Cohen, MD
Philadelphia, Pennsylvania Chicago, Illinois Professor
Departments of Radiology and Neurological
Syed T. Arshad, MD Jens Bracht, Dipl.-Phys. Surgery
Neuro-Intensivist Technical Director R&D, PD University of Washington School of Medicine
Department of Neurosurgery GMS mbH Seattle, Washington
Sacramento Medical Center/Kaiser Permanente Kiel, Germany
North Valley E. Sander Connolly, Jr., MD
Sacramento, California M. Ross Bullock, MD, PhD Bennett M. Stein Professor and Vice Chairman
Professor Department of Neurological Surgery
Ramani Balu, MD, PhD Department of Neurological Surgery Columbia University College of Physicians and
Fellow Director, Clinical Neurotrauma Surgeons;
Department of Neurology Department of Neurological Surgery Director, Cerebrovascular Research Laboratory
Division of Neurocritical Care University of Miami Miller School of Medicine Surgical Director, Neuro-Intensive Care Unit
Perelman School of Medicine at the University Miami, Florida Department of Neurological Surgery
of Pennsylvania Columbia University Medical Center/New
Philadelphia, Pennsylvania Andrew P. Carlson, MD York-Presbyterian
Department of Neurological Surgery New York, New York
Sarice L. Bassin, MD University of New Mexico School of Medicine
Assistant Professor Albuquerque, New Mexico Marek Czosnyka, PhD
Department of Neurology Reader in Brain Physics
Northwestern University Feinberg School of Emmanuel Carrera, MD Department of Clinical Neurosciences,
Medicine; Department of Neurology Neurosurgery Unit
Program Director, Neurocritical Care Fellowship Centre Hospitalier Universitaire Vaudois University of Cambridge/Addenbrooke’s
Division of Neurocritical Care (CHUV) Hospital
Northwestern Memorial Hospital Lausanne University Hospital Cambridge, United Kingdom
Chicago, Illinois Lausanne, Switzerland
John A. Detre, MD
David M. Benglis, Jr., MD Professor
Atlanta Brain and Spine Care Departments of Neurology and Radiology
Atlanta, Georgia Perelman School of Medicine at the University
of Pennsylvania
Philadelphia, Pennsylvania

ix
x Contributors

Martin E. Doerfler, MD Thomas Geeraerts, MD, PhD K.T. Henrik Huttunen, MD

Associate Professor Anesthesiology and Critical Care Department Clinical Assistant Professor of Anesthesiology
Departments of Medicine and Science Education University Hospital of Toulouse Department of Anesthesiology, Pharmacology,
Hofstra North Shore-LIJ School of Medicine at University Paul Sabatier and Therapeutics
Hofstra University Toulouse, France University of British Columbia Faculty of
Hempstead, New York; Medicine;
Vice President, Evidence Based Clinical Practice Vicente H. Gracias, MD Attending Anesthesiologist, Vancouver Acute
and Clinical Integration Professor Department of Anesthesia
North Shore-LIJ Health System Department of Surgery Division of Neuroanesthesia
Lake Success, New York Chief, Trauma/Surgical Critical Care Vancouver General Hospital
UMDNJ-Robert Wood Johnson Medical School; Vancouver, British Columbia, Canada
Guy M. Dugan, MD, FCCP Director, Level I Trauma Center
Director Robert Wood Johnson University Hospital Peter J. Kirkpatrick, MSc, FRCS(SN),
Department of Critical Care and Neurocritical New Brunswick, New Jersey FMedSci
Care Fellow of Medical Academy of Sciences
Alexian Brothers Medical Center David M. Greer, MD, MA, FCCM, FAHA Society of British Neurosurgeons (SBNS)
Elk Grove Village, Illinois Dr. Harry M. Zimmerman and Dr. Nicholas and Consultant Neurosurgeon
Viola Spinelli Professor and Vice Chairman Honorary University Lecturer
Richard P. Dutton, MD, MBA Director, Neurosciences Intensive Care Unit Department of Neurosurgery
Clinical Associate Neurology Residency Program Director Addenbrooke’s Hospital
Department of Anesthesia and Critical Care Director of Medical Studies Cambridge University Hospitals
University of Chicago; Department of Neurology Cambridge, United Kingdom
Executive Director Yale School of Medicine
American Quality Institute (AQI) New Haven, Connecticut Michel Kliot, MD
American Society of Anesthesiologists Professor of Clinical Neurosurgery
Park Ridge, Illinois Christiana E. Hall, MD, MS Department of Neurological Surgery
Associate Professor of Neurology and University of California, San Francisco School of
E. Wesley Ely, MD, MPH, FACP, FCCM Neurotherapeutics and Neurological Surgery Medicine;
Professor of Medicine and Critical Care Division of Neurocritical Care Director, Center for Management and Surgery of
Department of Medicine University of Texas Southwestern Peripheral Nerve Disorders
Center for Health Services Research Dallas, Texas San Francisco, California
Vanderbilt University School of Medicine;
Associate Director of Aging Research, VA J. Claude Hemphill III, MD, MAS W. Andrew Kofke, MD, MBA, FCCM
GRECC Professor of Clinical Neurology Professor
Veterans Affairs Tennessee Valley Healthcare Department of Neurology Departments of Anesthesiology and Critical
System University of California, San Francisco School of Care and Neurosurgery
Nashville, Tennessee Medicine; Perelman School of Medicine at the University
Director, Neurocritical Care of Pennsylvania;
Ronald G. Emerson, MD Department of Neurology Director, Neuroanesthesia
Adjunct Professor of Clinical Neurology San Francisco General Hospital Department of Anesthesiology and Critical Care
Department of Neurology San Francisco, California University of Pennsylvania Health System;
Columbia University Medical Center; Co-Director, Neurointensive Care Unit
Attending Neurologist Jiri Horak, MD Hospital of the University of Pennsylvania
Department of Neurology Assistant Professor of Clinical Anesthesiology Philadelphia, Pennsylvania
Hospital for Special Surgery and Critical Care
New York, New York Department of Anesthesiology and Critical Care Jaroslaw Krejza, MD, PhD
Perelman School of Medicine at the University Research Associate Professor
Per Enblad, MD, PhD of Pennsylvania Department of Radiology
Professor of Neurosurgery Philadelphia, Pennsylvania Perelman School of Medicine at the University
Department of Neuroscience of Pennsylvania
Section of Neurosurgery Peter Horn, MD Philadelphia, Pennsylvania;
Uppsala University Associate Professor of Neurosurgery Al-Imam Muhammad Ibn Saud Islamic
Uppsala University Hospital Department of Neurosurgery University
Uppsala, Sweden Charité Universitätsmedizin Berlin Riyadh, Saudi Arabia
Berlin, Germany
Anthony A. Figaji, MD, FCS, PhD Monisha A. Kumar, MD
Professor David A. Horowitz, MD Assistant Professor
Division of Neurosurgery Assistant Professor of Clinical Medicine Departments of Neurology, Neurosurgery, and
University of Cape Town; Perelman School of Medicine at the University Anesthesiology and Critical Care
Head of Pediatric Neurosurgery of Pennsylvania; Perelman School of Medicine at the University
Red Cross Children’s Hospital Associate Chief Medical Officer of Pennsylvania;
Cape Town, Western Cape, South Africa University of Pennsylvania Health System Director, Neurocritical Care Fellowship Program
Philadelphia, Pennsylvania Hospital of the University of Pennsylvania
Damien Galanaud, MD, PhD Philadelphia, Pennsylvania
CNRS UPR 640 LENA
Université Pierre et Marie Curie (Paris VI);
Neuroradiology
Pitié Salêtrière Hospital
Paris, France
 Contributors xi

Arthur M. Lam, MD, FRCPC David K. Menon, MD, PhD, FRCP, FRCA, DaiWai M. Olson, PhD, RN
Medical Director of Neuroanesthesia and FFICM, FMedSci Assistant Professor
Neurocritical Care Professor and Head Department of Medicine
Swedish Neuroscience Institute Division of Anaesthesia Division of Neurology
Swedish Medical Center; University of Cambridge/Addenbrooke’s Duke University School of Medicine
Clinical Professor of Anesthesiology and Pain Hospital; Durham, North Carolina
Medicine Honorary Consultant
University of Washington Perioperative Care Pratik P. Pandharipande, MD, MSCI
Member, Physician Anesthesia Services Addenbrooke’s Hospital; Associate Professor
Seattle, Washington Co-Chair, Acute Brain Injury Program Department of Anesthesiology
Wolfson Brain Imaging Centre Division of Critical Care and Perioperative
Peter D. le Roux, MD, FACS University of Cambridge Medicine
Associate Professor of Neurosurgery Cambridge, United Kingdom Vanderbilt University School of Medicine
Perelman School of Medicine at the University Nashville, Tennessee
of Pennsylvania; Asako Miyakoshi, MD
Department of Neurosurgery Assistant Professor Jose L. Pascual, MD, PhD, FRCS(C), FACS
Pennsylvania Hospital Department of Radiology Assistant Professor
Philadelphia, Pennsylvania Division of Neuroradiology Department of Surgery
University of Washington School of Medicine Division of Traumatology, Surgical Critical Care,
Joshua M. Levine, MD Seattle, Washington and Emergency Surgery
Assistant Professor Perelman School of Medicine at the University
Departments of Neurology, Neurosurgery, and Richard S. Moberg, MSE of Pennsylvania;
Anesthesiology and Critical Care President Attending Surgeon
Perelman School of Medicine at the University Moberg Research, Inc. Department of Surgery
of Pennsylvania; Ambler, Pennsylvania Hospital of the University of Pennsylvania
Co-Director, Neurointensive Care Unit Philadelphia, Philadelphia
Hospital of the University of Pennsylvania Pierre D. Mourad, PhD
Philadelphia, Pennsylvania Associate Professor Aashish R. Patel, DO
Department of Neurological Surgery Fellow in Neurocritical Care
Geoffrey T. Manley, MD, PhD Principal Physicist Department of Neurological Surgery
Professor and Vice Chairman Applied Physics Laboratory University of Texas Southwestern
Co-Director, Brain and Spinal Injury Center University of Washington Dallas, Texas
Department of Neurological Surgery Seattle, Washington
University of California, San Francisco School of Frederik A. Pennings, MD, PhD
Medicine; Barnett R. Nathan, MD Assistant Professor
Chief of Neurosurgery Associate Professor Department of Neurosurgery
San Francisco General Hospital Departments of Neurology and Internal University of Massachusetts Medical School
San Francisco, California Medicine Worcester, Massachusetts
University of Virginia
Basil F. Matta, MB, MA, BCh, FRCA Charlottesville, Virginia Ian Piper, PhD
Associate Lecturer Clinical Scientist
Department of Medicine Patrick J. Neligan, MD Clinical Physics
Division of Anaesthesia Senior Clinical Lecturer of Anaesthesia and University of Glasgow;
University of Cambridge/Addenbrooke’s Intensive Care Brain-IT Group Coordinator, Intensive Care
Hospital; Galway University Hospitals Monitoring
Divisional Director and Associate Medical National University of Ireland, Galway Department of Clinical Physics
Director Galway, Ireland Southern General Hospital Trust
Emergency and Perioperative Care Glasgow, United Kingdom
Cambridge University Foundation Trust Anoma Nellore, MD
Hospitals Fellow Amit Prakash, MBBS, MD, FRCA, EDIC
Cambridge, United Kingdom Department of Medicine Consultant, Department of Anaesthesia and
Division of Infectious Disease Intensive Care
Jonathan McEwen, MD Massachusetts General Hospital Addenbrooke’s Hospital
Clinical Assistant Professor of Anesthesiology Harvard Medical School Cambridge University Hospital Foundation
Department of Anesthesiology, Pharmacology, Boston, Massachusetts Trust
and Therapeutics Cambridge, United Kingdom
University of British Columbia Faculty of Mauro Oddo, MD
Medicine; Staff Physician J. Javier Provencio, MD, FCCM
Attending Anesthesiologist, Vancouver Acute Department of Intensive Care Medicine Associate Professor
Department of Anesthesia, Division of CHUV-University Hospital Departments of Neurology, Neurological
Neuroanesthesia Faculty of Biology and Medicine University of Surgery, and Neurosciences
Vancouver General Hospital Lausanne Cleveland Clinic Lerner College of Medicine of
Vancouver, British Columbia, Canada Lausanne University Hospital Case Western Reserve University;
Lausanne, Switzerland Director, Neurocritical Care Fellowship Program
Cleveland Clinic
Cleveland, Ohio
xii Contributors

Louis Puybasset, MD, PhD J. Michael Schmidt, PhD, MSc Martin Smith, MBBS, FRCA, FFICM
Département d’Anesthésie-Réanimation Assistant Professor of Clinical Neurophysiology Consultant and Honorary Professor in
Université Pierre et Marie Curie (Paris VI); in Neurology Neurocritical Care
Neurosurgical Intensive Care Unit Department of Neurology The National Hospital for Neurology and
Pitié Salêtrière Hospital Columbia University College of Physicians and Neurosurgery
Paris, France Surgeons; University College London Hospitals
Director London, United Kingdom
Rohan Ramakrishna, MD Neuro-ICU Neuromonitoring and Informatics
Resident Columbia University Medical Center Marco D. Sorani, PhD
Department of Neurological Surgery New York, New York Adjunct Assistant Professor
University of Washington Medical Center Department of Neurological Surgery
Seattle, Washington Sarah E. Schmitt, MD University of California, San Francisco
Assistant Professor of Clinical Neurology San Francisco, California
Mahbub Rashid, PhD Department of Neurology
Professor Perelman School of Medicine at the University Alejandro M. Spiotta, MD
University of Kansas School of Architecture, of Pennsylvania; Resident
Design and Planning Director, Electroencephalography Laboratory Department of Neurosurgery
Lawrence, Kansas Department of Neurology Cleveland Clinic
Hospital of the University of Pennsylvania Cleveland, Ohio
Gerald P. Roston, PhD, PE Philadelphia, Pennsylvania
Technology Consultant and Managing Partner John J. Stern, MD
Pair of Docs Consulting, LLC Patricia D. Scripko, MD Clinical Professor
Saline, Michigan Resident Department of Medicine
Department of Neurology Perelman School of Medicine at the University
Stuart Russell, PhD Massachusetts General Hospital of Pennsylvania;
Professor of Computer Science Brigham and Women’s Hospital Chief, Division of Infectious Diseases
Michael H. Smith and Lotfi A. Zadeh Chair in Boston, Massachusetts Department of Medicine
Engineering Pennsylvania Hospital
Computer Science Division †John M. Sewell, BSEE Philadelphia, Pennsylvania
University of California, Berkeley Chief Engineer
Berkeley, California; Active Signal Technologies, Inc. Nino Stocchetti, MD
Adjunct Professor Linthicum, Maryland Professor of Anesthesia and Intensive Care
Department of Neurological Surgery Terapia Intensiva Neuroscienze
University of California, San Francisco Robert G. Siman, PhD Fondazione IRCCS Cà Granda
San Francisco, California Research Professor University of Milan
Department of Neurosurgery Milan, Italy
Owen B. Samuels, MD Perelman School of Medicine at the University
Associate Professor of Neurosurgery and of Pennsylvania Jose I. Suarez, MD
Neurology Philadelphia, Pennsylvania Professor
Emory University School of Medicine; Departments of Neurology and Neurosurgery
Director, Division of Neuroscience Critical Care Carrie A. Sims, MD, FACS Baylor College of Medicine;
Emory Healthcare Assistant Professor Director
Atlanta, Georgia Department of Surgery Vascular Neurology and Neurocritical Care
Division of Traumatology, Surgical Critical Care, Baylor College of Medicine
Matthew R. Sanborn, MD and Emergency Surgery Houston, Texas
Resident Perelman School of Medicine at the University
Department of Neurosurgery of Pennsylvania Farzana Tariq, MD
Perelman School of Medicine at the University Philadelphia, Pennsylvania Cerebrovascular and Skull Base Fellow
of Pennsylvania Department of Neurological Surgery
Philadelphia, Pennsylvania Richard O. Sinnott, PhD University of Washington
Director, eResearch Seattle, Washington
Bernhard Schmidt, PhD University of Melbourne
Department of Neurology Melbourne, Australia Kyla P. Terhune, MD
Chemnitz Medical Centre Assistant Professor of Surgery and
Chemnitz, Germany Alan Siu Anesthesiology
Resident Division of General Surgery
Eric Albert Schmidt, MD, PhD Department of Neurological Surgery Vanderbilt University School of Medicine
Department of Neurosurgery The George Washington University Medical Nashville, Tennessee
Hôpital Purpan Center
Toulouse, France Washington, District of Columbia Brett Trimble, BSME
Director, Advanced Technology
Integra LifeSciences Corporation
†Deceased San Diego, California
 Contributors xiii

David K. Vawdrey, PhD Brandon von Tobel, MD, MBE Elisa R. Zanier, MD
Assistant Professor of Clinical Biomedical Vice President of Finance and Operations Department of Neuroscience
Informatics ImaCor, Inc. Instituto Mario Negri
Department of Biomedical Informatics New York, New York Milan, Italy
Columbia University College of Physicians and
Surgeons Howard Yonas, MD Craig Zimring, PhD
New York, New York Professor and Chairman Professor of Architecture and Psychology
Department of Neurosurgery Colleges of Architecture and Psychology
Paul M. Vespa, MD University of New Mexico School of Medicine Georgia Institute of Technology
Professor of Neurosurgery and Neurology Albuquerque, New Mexico Atlanta, Georgia
Department of Neurosurgery
David Geffen School of Medicine at UCLA; Brad E. Zacharia, MD
Director Resident
Neurocritical Care Department of Neurological Surgery
Ronald Reagan UCLA Medical Center Columbia University Medical Center/New
Los Angeles, California York-Presbyterian
New York, New York
Abbreviations
NOTE: Abbreviations may have more than one meaning, depending on their context.

3-H hypertension, hemodilution, and hypervolemia ASTM American Society for Testing and Materials
AACN American Association of Critical Care Nurses AT antithrombin
AAN American Academy of Neurology ATC automatic tube compensation
AARP American Association of Retired Persons ATN acute tubular necrosis
ABA American Bar Association ATP adenosine triphosphate
ABM acute bacterial meningitis ATS American Thoracic Society
ABP arterial blood pressure AU arbitrary units
ACA anterior cerebral artery AV audiovisual
ACE angiotensin converting enzyme AVDO2 arteriovenous difference in oxygen
ACEI/ARBs angiotensin-converting enzyme inhibitor/ AVM arteriovenous malformation
angiotensin-receptor blocker BA basilar artery
ACGME Accreditation Council for Graduate Medical Education BAEP brainstem auditory evoked potential
AChE acetylcholinesterase BAM brain acoustic monitor
ACCP American College of Chest Physicians BANN Bayesian Artificial Neural Network
ACoA anterior communicating artery BBB blood-brain barrier
ACS abdominal compartment syndrome BFV blood flow velocity
ACT activated clotting time BG blood glucose
ACTH adrenocorticotrophic hormone BHI breath holding index
ACV assist control ventilation BIS bispectral index
AD axial diffusivity BMI body mass index
ADC apparent diffusion coefficient BMR basal metabolic rate
ADH antidiuretic hormone BOLD blood oxygen level dependent
ADL activities of daily living BOOST Brain Oxygen and Outcome Study in Traumatic Brain Injury
ADNI Alzheimer’s Disease Neuroimaging Database BP blood pressure
ADP adenosine diphosphate BPI bactericidal permeability-increasing protein
ADQI Acute Dialysis Quality Initiative BrainIT brain monitoring with information technology
ADR alpha/delta ratio BSI bloodstream infection
ADT admission/discharge/transfer BSM bedside monitors
AED antiepileptic drug BT brain temperature
aEEG amplitude-integrated electroencephalography BTO balloon test occlusion
AEP auditory evoked potentials BUN blood urea nitrogen
AF atrial fibrillation CA cerebral autoregulation (Chapters 30, 46)
AG anion gap CA cardiac arrest (Chapter 25)
AHA/ASA American Heart Association/American Stroke Association CA-BSI catheter-associated bloodstream infection
AI artificial intelligence CAD coronary artery disease
AIS acute ischemic stroke CAM-ICU Confusion Assessment Method for the ICU
AKI acute kidney injury CAP College of American Pathologists
AKIN Acute Kidney Injury Network CAR cerebral arterial resistance
ALF acute liver failure CAS carotid angioplasty and stenting
ALFSG Acute Liver Failure Study Group CASL continuous arterial spin labeling
ALI acute lung injury CBF cerebral blood flow
AMA American Medical Association CBFV cerebral blood flow velocity
AMID active implantable medical device CBV cerebral blood volume
ANH artificial nutrition and hydration CCAT Computerized Cognitive Assessment Tool
ANN artificial neural network CCO continuous cardiac output
APACHE Acute Physiology and Chronic Health Evaluation CCT central conduction time
aPL antiphospholipid antibodies Ccw compliance of the chest wall
APN advanced practice nurse CDC Centers for Disease Control and Prevention
APP abdominal perfusion pressure CDSA color density spectral array
aPTT activated partial thromboplastin time CEA carotid endarterectomy
ARAS ascending reticular activating system cEEG continuous electroencephalography
ARC absolute reticulocyte count CES cholesterl emboli syndrome
ARDS acute respiratory distress syndrome CEUs continuing education units
ARi (or ARI) autoregulation index CFM cerebral function monitoring
ASA American Society of Anesthesiology C-FMZ C-flumazenil
aSAH aneurysmal subarachnoid hemorrhage Cho choline
ASIA American Spinal Injury Association CHr reticulocyte hemoglobin content
ASL arterial spin labeling CI coagulation index

xv
xvi Abbreviations

CINMA critical illness neuromuscular abnormalities DHHS Department of Health and Human Services
CIPM critical illness polyneuromyopathy DI diabetes insipidus
CIPNM critical illness polyneuropathy and myopathy DIC disseminated intravascular coagulation
CIRCI critical illness related corticosteroid insufficiency DIND delayed ischemic neurologic deficit
CK creatine kinase DIT drug-induced thrombocytopenia
Cl compliance of the lung DITP drug-induced immune thrombocytopenia
CLIA 88 Clinical Laboratory Improvements Amendments of 1988 dIVC inferior vena cava diameter
CLAB central line-associated bacteraemia DLCO diffusing capacity of the lung for carbon monoxide
CLABSI central line-associated bloodstream infection DMN default mode network
CMAP compound muscle action potentials DO2 oxygen delivery
CMO comfort measures only DoD Department of Defense
CMRO2 cerebral metabolic rate of oxygen DoE Department of Energy
CMRGluc cerebral metabolic rate of glucose DRG diagnostic related group
CMS Centers for Medicare and Medicaid Services DRS Disability Rating Scale
CMV cytomegalovirus DS Down syndrome
CNS central nervous system DSA digital subtraction angiography
CO cardiac output DSM Diagnostic and Statistical Manual of Mental Disorders
CO2 carbon dioxide DSP digital signal processing
COGIF Consensus on Grading Intracranial Flow DTI diffusion tensor imaging
COI cerebral oxygenation index DUS duplex ultrasonography
COM communication DV data validation
COMBI Center for Outcome Measurement in Brain Injury DVT deep vein thrombosis
COMPACCS Committee on Manpower for Pulmonary and Critical DWI diffusion-weighted imaging
Care Societies EAA excitatory amino acids
COPD chronic obstructive pulmonary disease EBM evidence-based medicine
COx cerebral oximetry index EBNP evidence-based nursing practice
CPAP continuous positive airway pressure EBP evidence-based practice
CPOE computerized order entry EC-IC extracranial to intracranial
CPP cerebral perfusion pressure ECA external carotid artery
CPR cardiopulmonary resuscitation ECCO Essentials of Critical Care Orientation
CPSE complex partial status epilepticus ECF extracellular fluid
CPT current procedural terminology ECoG electrocorticogram
Cr creatine ED emergency department
CRH corticotropin-releasing hormone EDC extended differential count
CRM crew/crisis resource management EDH epidural hematoma
CRMP collapsin response mediator protein EDM esophageal Doppler monitor
CRP C-reactive protein EDTA ethylene diamine tetra acetate
CRRT continuous renal replacement therapy EEG electroencephalography; electroencephalogram
CRS Coma Recovery Scale EF ejection fraction
CRS-R Coma Recovery Scale–Revised E-GOS Extended Glasgow Outcome Scale
Crs compliance of respiratory system EHR electronic health record
CSA cross-sectional area EIT electrical impedance tomography
CSD cortical spreading depression EKG electrocardiogram
CSE convulsive status epilepticus ELISA enzyme-linked immunological sample assay
CSF cerebrospinal fluid EMG electromyogram
CSW cerebral salt wasting EMI electromagnetic interference
CT computed tomography EMR electronic medical record
CTA computed tomography angiography EMS emergency medical services
CTP computed tomography perfusion (Chapters 13, 26) EN enteral nutrition
CTP Child-Turcotte-Pugh (Chapter 23) eNAA extracellular N-acetyl aspartate
CTT central conduction time EOG electrooculogram
CTV cerebral venous thrombosis EP evoked potential
CVC central venous catheter EPIC extended prevalence of infection in intensive care
CVP central venous pressure EPL estimated percent lysis
CVR cerebrovascular resistance EPO erythropoietin
CVT cerebral venous thrombosis EPOR erythropoietin receptor
CVVH continuous veno-venous hemofiltration ESA erythropoiesis-stimulating agents
CXR chest x-ray ESICM European Society of Intensive Care Medicine
D diameter of conduit ESO European Stroke Organization
DAI diffuse axonal injury ESRD end-stage renal disease
dARi dynamic autoregulation index ET endotracheal tube
DBN dynamic Bayesian network etCO2 end-tidal carbon dioxide
DBP diastolic blood pressure ETF Emerging Technology Fund
DBS deep brain stimulator EU European Union
DC decompressive craniectomy EVD external ventricular drain
DCI delayed cerebral ischemia EVLWI extravascular lung water index
DCS diffuse correlation spectroscopy FA fractional anisotropy
DHCA deep hypothermic circulatory arrest FC Foley catheter
Other documents randomly have
different content
 In illustration of this subject, I would first direct attention to the
beautiful fossil, figured Lign. 188, fig. 1, which was discovered in the
Chalk near Lewes, and is figured, of the natural size, Foss. South D.
tab. xxxix. This ray, or spine, belongs to one of the Cestraciont fishes
(Ptychodus), whose teeth are so abundant in the Chalk, and will
presently be described. It is composed of fourteen thick, flat, osseous
rods, or strands, intimately united together, with longitudinal furrows
or sutures on the surface. The anterior margin is embossed, and the
projections form on the sides wide, rounded ribs, and transverse
depressions. Towards the base of the posterior part, there are large
osseous fibres inserted vertically and obliquely, which appear to have
been processes of attachment. The rods, or plates, are parallel With
the posterior margin, and each terminates in a rounded extremity, or
boss, on the front edge of the spine. This ray is wider at its base than
at the superior part the anterior margin is oblique, and the posterior
straight. The surface, where entire, is covered with a dense osseous
substance, which is finely engrailed.[515]
[515] This specimen is figured in Poiss. Foss.; but it is
represented too short, from the two portions being drawn as if they
were connected, without any interval between them, as in Lign.
188. It is in the British Museum. See Petrifactions, p. 450.

Lign. 188. Dorsal Rays of Sharks. Sussex.

Fig. 1.— Ptychodus spectabilis. 1/5 nat. Chalk. Lewes.
 1a. Portion of a Dorsal Ray, with oblique serrated
— sutures. Chalk. Lewes.
2.— Ptychodus Gibberulus. 1/5 nat. Chalk. Lewes.
3.— Hybodus subcarinatus. nat. Weald. Tilgate Forest.

In 1850 I discovered in the Plastic Clay of Castle Hill Newhaven, a

dorsal fin of Ptychodus, with eight vertebræ. A nearly entire fin-ray of
this species, three feet in length, has recently been discovered by Mr.
Charles Potter, of Lewes, in the Chalk near that town. The remains of
another ray, of equal proportions, were found near it; and these dorsal
spines might have belonged to the same individual, for there are no
reasons to forbid the supposition that the Ptychodus had two dorsal
fin-rays. The length of these spines necessarily indicates a very large
fish.
A smaller species of Ichthyodorulite, also found in the Lewes Chalk,
is distinguished from P. spectabilis by its osseous plates contracting
towards their extremities, and terminating more suddenly on the front
margin, producing gibbosities less acute and more distant than in P.
spectabilis; this species is named P. gibberulus: see Lign. 188, fig. 2.
[516]

[516] This fossil is figured of the natural size, Foss. South D. pl.
xl. fig. 3.

The bony plates of these fins are occasionally found lying in

irregular groups in the Chalk, as if the fin had partially decomposed
and the plates separated. In one example, the rays are split asunder
by a piece of bone, apparently a portion of a long pointed tooth, firmly
impacted between them; as if the fish had been seized by some
enemy, and had escaped, with the tooth of its adversary in its fin. Very
fine specimens have been found at Charing, Kent, by W. Harris, Esq.
F.G.S.
In the fragment of an Ichthyodorulite from the Lewes Chalk, a
remarkable structure is displayed; the osseous plates are united
laterally by smooth, longitudinal lines, as in those above described; but
they are also traversed by numerous oblique, finely-serrated sutures.
Lign. 188, fig. 1a.
 The Chalk contains rays of other species of Ptychodus, as well as of
some allied genera. Of these, the most remarkable are smooth,
arched, pointed spines, having a shallow posterior groove, with an
enamelled surface, marked with fine longitudinal striæ, and frequent,
parallel, oblique lines. These, according to Sir P. Egerton, belong to a
true Cestracion (see p. 584): they were first figured and described by
me (Foss. South D. tab. xxxiii. fig. 5) as belonging to the Acanthias
major, and were subsequently assigned to the genus Spinax by Prof.
Agassiz (Poiss. Foss. iii. p. 62).
It may be necessary to remark, that the fins first described have
been referred to the fishes which yielded the large grooved teeth so
common in the Chalk (see Pl. VI fig. 2) in consequence of their affinity
to existing species, which have similar fins and teeth; and from the
circumstance that the Sharks of the genus Lamna, whose teeth also
abound in the Chalk, have no dorsal rays of this kind; still the proof of
identity remains to be discovered. In one specimen only have I
observed indications of any other part of the skeleton; it is a spine of
Acanthias major, the base of which rests on several dorsal vertebræ
(Foss. South D. tab. xxxiii.).
Hybodus subcarinatus. Lign. 188, fig. 3.—The fishes of another extinct
genus of Sharks, termed Hybodus, from the gibbous form of the teeth,
were also provided with dorsal spines, which may be readily
distinguished from the preceding. These Ichthyodorulites are generally
long, slightly arched, and terminate in a point at the extremity; the
base, which was implanted in the flesh, is deeply grooved, and much
prolonged, being sometimes equal to one-third of the entire length.
The surface is marked with strong longitudinal ridges, parallel with the
anterior margin which is rounded and laterally compressed. The
posterior edge, which is more or less flat, has, towards the base two
rows of sharp arched teeth, which gradually approach ’each other, and
blend into one line on the upper part of the ray There are numerous
species of this genus in the Oolite and Lias. I have found one species
in the Chalk and a few in the Wealden. The small Ray figured Lign. 188
fig. 3, is from Tilgate Forest, and displays the usual characters of these
fossils. From specimens discovered in the Lias, associated with the
teeth, it appears that the Hybodus had two dorsal fins, each furnished
with rays, as in the recent Dog-fish.
The microscopic structure of these rays is stated by M. Agassiz to
resemble that of the teeth: in some there is a pulp cavity, which
occupies the centre of the spine, and is surrounded by dentine, in
which the calcigerous tubes radiate direct to the surface; the external
enamel is a layer of dentine, in which the medullary canals are
wanting.
In the strata below the Lias there are numerous Ichthyodorulites,
some of a large size, belonging chiefly to the Cestracion family, and of
extinct species, not observed in more recent deposits. Thus there are
several species of dorsal rays (named Onchus, from their hooked form,)
that are wide at the base, and bent backwards, with the posterior
margin destitute of teeth, in the Carboniferous, Devonian, and Silurian
formations; also immense compressed spines, having small teeth on
the posterior margin, and the surface covered with longitudinal striæ,
and finely toothed, transversely; hence termed Ctenacanthus, or
pectinated-spine (Murch. Sil. Syst. p. 596).
The fossil spine, named Orthacanthus (Poiss. Foss. vol. iii. pl. xlv.),
and found in the Coal of Manchester, has been discovered in connexion
with the body of the fish to which it belonged in the Carboniferous
deposits of Ruppersdorf in Bohemia (Geol. Journal, vol. v. part ii. p.
23).
Some Ichthyodorulites have the surface richly ornamented with
stellular tubercles, and are termed Asteracanthus, or starry-spine; there
are very large fin-rays of this kind in the Wealden, Purbeck, Oolites,
and Lias.[517]
[517] For particular information on Ichthyodorulites, consult
Poiss. Foss. tom. iii. chap. i. About seventy species are enumerated.

The Ichthyodorulites of the Rays have no cavity like those of the

Sharks, and are of a depressed form, and more or less flattened; they
are armed with teeth along their exterior margins, and not on the
posterior edge, as in the latter family.
 Fossil Teeth of Fishes.—From the durable nature and striking
appearance of many of the fossil teeth of fishes, and their prodigious
numbers in some deposits, they are familiar objects to the collector. By
far the largest proportion of the detached teeth belongs to various
species and genera of that most numerous, and widely distributed
family of voracious fishes, the Sharks. In the Tertiary strata teeth of this
kind occur of a very large size; in the Chalk many species abound,
particularly of the lanceolate and compressed forms, and of the
rugous, mammillated, palatal teeth, commonly termed palates. As we
pass to the more ancient formations, teeth of different forms prevail;
and those which approach the recent types are either very rare or
altogether absent. We will select some examples of the different
genera in illustration of this subject; the previous observations on the
form and structure of the recent teeth render but few introductory
remarks necessary.
Fossil Teeth of Sharks.—The fishes of the Shark and Ray families
belong to the Placoid order; the scales in the former consist of
enamelled plates and tubercles, forming a shagreen surface; and in
the dermal integument of the latter they appear as spines and bosses,
irregularly disposed.
Notwithstanding the diversity in appearance of the teeth of Sharks,
they all possess one essential character of structure, namely, a base,
or osseous root of variable form, which is implanted in the
integuments; and a crown, or external portion, which projects into the
mouth, is covered with enamel or compact dentine, and assumes
numerous modifications, by which the fossil genera are characterized.
These teeth are never imbedded in sockets, nor united to the dentary
margins of the jaws; they only adhere to the integuments of the
mouth, and the covering of the maxillæ; they possess, in most of the
Sharks, great mobility. They are generally disposed in rows; the
anterior ones, being first used, fall out, and are replaced by those on
the inner series. New teeth are also continually formed behind those in
use, and advance successively towards the anterior rows as the latter
are shed, and in their turn occupy the front rank. (See Cyclop. Anat.
Art. Teeth.) An examination of the fossil and recent teeth of Sharks and
Rays proves that the prevailing existing generic types have but few, if
any, representatives in the fossils, except in those which belong to the
Tertiary and Cretaceous formations; while the genera that appear
isolated, as it were, in the present seas have numerous analogues in
the Secondary strata.
The fossil teeth of this family may be divided into two grand
divisions; namely, those which are more or less of a polygonal,
obtusely conical, or depressed form, having a tesselated arrangement
in the mouth; and those of a triangular, lanceolate shape, with cutting,
or serrated edges, disposed in a series of rows on the jaws. The teeth
of the first group (Cestracionidæ) have most analogy to those of the
living genus Cestracion (Port-Jackson Shark); the second (Sgualidæ) to
the Sharks, commonly so called.
The Cestracion is the only living representative of a family of
squaloid fishes of a peculiar type, whose remains occur in almost the
earliest fossiliferous deposits; it inhabits the seas of New Holland and
the southern coasts of China. The jaws of the Cestracion are relatively
very large, and are armed with numerous rows of teeth, essentially of
two kinds; those situated anteriorly, or towards the front of the mouth,
being adapted for seizing and retaining the food, and the posterior
ones for crushing and bruising. The prehensile teeth are sharp,
angular, and pointed: the others are obtuse, polygonal, enamelled, and
disposed in oblique rows along the margins and inner surface of both
jaws; there are sometimes sixty in each jaw (see Bd. ii. pl. xxvii 11. fig.
A). Fossil teeth of this type are exceedingly numerous in the Chalk,
Lias, &c. but are very seldom found in juxtaposition; the
decomposition of the cartilaginous integuments in which they are
imbedded, having, in most examples, occasioned their displacement
and dispersion; specimens, however, are occasionally discovered, in
which numerous teeth, of various sizes, are disposed in mosaic, in
their natural relative positions.
The extinct forms of this family (Cestracionidæ) are known almost
only by their teeth; and according to the shape, structure, and
sculpture of these organs, M. Agassiz has arranged them into several
genera. They occur in most of the fossiliferous deposits.
 Cestracion canaliculatus.—The teeth of a fish belonging to the
existing genus have been discovered in the Chalk of Kent; they are
figured and described by Sir P. Egerton in the beautiful work by Mr.
Dixon.[518] This unique specimen consists of a group of thirteen
posterior molar and three or four detached prehensile anterior teeth,
imbedded in a block of chalk about two inches square. The chief
distinction from the teeth of the recent Cestracion is in the presence of
a large medullary canal which traverses the base of each tooth: hence
the specific name.
[518] Dixon’s Fossils of Sussex, &c. p. 365, tab. xxxii. fig. 8.
From the t examination of a specimen lately found at Lewes, Sir P.
Egerton has been led to assign to this species the spine formerly
described as Spinax major.

Acrodus (ridge-tooth) nobilis. Lign. 189, fig. 4, Ly. p. 275, fig. 307.—
In the Lias and Oolite, oblong enamelled teeth, having the surface of
the crown covered with fine radiating grooves and striæ, are well
known to collectors, in many parts of England, by the name of fossil
leeches, from a fancied resemblance to a contracted leech. They
belong to an extinct genus of Cestracionts, named Acrodus by M.
Agassiz. The crown of the tooth is enamelled, and covered with
transverse grooves, which diverge from a longitudinal furrow; the base
is in the form of a parallelogram inclined on its inner side. These teeth
were inserted along the jaws in oblique series, their longitudinal
direction corresponding with that of the bones which supported them;
in their natural position, the extremity of a hinder tooth was enclosed
between the two next anterior teeth. A beautiful group is figured Bd. ii.
pl. xxviie.[519]
[519] The microscopical structure of the teeth of Acrodus is well
shown in the "Odontography," pl. xiv. xv., and beautifully illustrates
the relation of dentine to bone.
 Lign. 189. Fossil Teeth of Sharks.
Fig. 1. Ptychodus Mortoni. (G. A. M.) Cret.
— New Jersey.
2.
Psammodus cinctus. (Ag.) Mt. L. Bristol.
—
3.
Orodus cinctus. (Ag.) Mt. L. Bristol.
—
4.
Acrodus nobilis. Lias. Lyme Regis.
—

Ptychodus (wrinkle-tooth). Pl. VI. fig. 2; Lign. 189, and Lign. 191.—
The palatal teeth, which occur more or less abundantly in almost every
chalk-pit, and are known by the name of "palates," belong to several
species of the genus Ptychodus. A very common form is figured Pl. VI.
fig. 2; and microscopic views of vertical and transverse sections, as
seen by transmitted light, are shown in figs. 2b, 2c. Groups of these
teeth, somewhat naturally arranged, and varying in size and form
according to the situations they occupied in the jaws, are occasionally
found: one specimen in the British Museum, and formerly in my
collection, contains more than 120 teeth. In general they occur in a
very perfect state, with the osseous base and enamelled crown entire.
The dorsal rays or spines previously described (p. 577), are sometimes
found with the teeth, and belong to fishes of the same genus.
These teeth are of an angular form, and more or less square, the
crown is wider than the root, which is obtuse, truncated, and
depressed in the centre; the enamelled part of the tooth is expanded
at the edges, and forms in the centre a flattened or slightly convex
mammillary projection, which is traversed by large, acute, transverse,
parallel ridges. The borders are granulated, and the sides of the
projection marked with deep vertical plicæ or folds; this description
particularly applies to the species named P. polygurus, figured in Plate
VI. Dr. Buckland has represented a fine group of these teeth, Bd. ii. pl.
xxvi′. Another common species (P. decurrens) is distinguished from the
former by the connexion between the large furrows on the crown and
the granulations on the expanded border, which diverge from the outer
edge of the large folds to the margin of the enamel.
The microscopic structure of these teeth presents the same
congeries of medullary and calcigerous tubes as those of the recent
Cestracion: see Plate VI. figs. 2b, 2c.
The teeth of a species of Ptychodus occur in the arenaceous strata
of the Chalk-formation in New Jersey, which possess the essential
characters of the European types, but differ from them in their
configuration; the only specimen I have seen is figured Lign. 189, fig.
1; it was presented to me by Dr. Morton. The enamelled crown forms a
conical projection, traversed by large inosculating ridges, which radiate
from the summit towards the margin.[520]
[520] I have named it P. Mortoni, in honour of my distinguished
friend, the eminent American naturalist and physician, Dr. George
Morton, of Philadelphia, by whom it was discovered.

Psammodus[521] (sandy-tooth). Pl. VI. fig. 1; Lign. 128, fig. 2.—To

this genus are referred the fossil teeth of the extinct Cestracionts,
which have the crown formed of small vertical tubes, with the grinding
surface more or less smooth, and presenting only a punctated or
sandy appearance.[522] These teeth are generally flat or slightly
convex, and of a square or oblong form; the base is osseous, and as
large as the crown. Two species are figured, Lign. 189, fig. 2, and Pl.
VI. fig. 1a. A magnified vertical section of the crown, displaying the
medullary canals and radiating calcigerous tubes, is represented Pl. VI.
fig. 1b, and a transverse section, fig. 1c; they are thin slices of a tooth,
P. porosus, from the Black Rock (Mountain Limestone), near Clifton,
viewed by transmitted light. The large, flat, quadrilateral, oblong teeth
that abound in the Stonesfield Slate belong to the Strophodus magnus.
[521] See Odontography, pl. xviii. xix.
[522] Ibid. vol. i. p. 59.

There are several kinds of fossil teeth which possess the same
essential structure as those of Psammodus, but differ in their external
characters; these are referred to other genera by M. Agassiz. Thus
Orodus, Lign. 189, fig. 3, comprises those elongated teeth in which the
centre of the crown forms an obtuse transverse cone, traversed by a
ridge from which oblique furrows diverge transversely towards the
circumference. Similar teeth, but with a smooth, obtusely conical
crown, are referred to the genus Helodus. Those with the crown
compressed and elevated, with a sharp edge, and with the base
surrounded by concentric folds, constitute the type of Chomatodes. A
similar crown, but subdivided by deep transverse ridges into
dentations, characterises the genus Ctenoptychius.
Ceratodus (horn-tooth) emarginatus. Lign. 194, fig. 1.—Very curious
dental organs, possessing a structure analogous to that of the teeth of
Psammodus, are found in the Bone-bed of the Lias; they consist of
consolidated plates instead of separate teeth; there was probably but
one plate on each side the jaws. The upper margin is generally
undulated, and more or less worn by use. These dental plates are
composed of two distinct layers; the lowermost portion, or root, is an
osseous, reticulated tissue, as in cartilaginous fishes in general; and
the upper consists of dentine, with minute parallel vertical tubes, as in
Psammodus; these tubes are a continuation of the medullary tissue of
the osseous root.
One species occurs in the Great Oolite at Stonesfield, and very
many forms abound in the Bone-bed at Aust Cliff, near Westbury on
Avon: and in the Trias (bone-bed) of Germany the teeth of several
species of Ceratodus are very abundant.
 The fishes to which these fossil teeth, referred to Ceratodus,
belonged were most probably Cestracionts; the ray-spine known as
Nemacanthus is provisionally assigned to them.
Edaphodon. Lign. 190 and Lign. 191, Ly. p. 276, fig. 309.—The
Chimæroid fishes, though formerly placed with the Plagiostomes
(Sharks and Bays), constitute a distinct group, of which there are but
two recent genera, though several occur in a fossil state. Their dental
organs are very peculiar. Their mandibles are furnished with two or
more pairs of oblong teeth, composed of long hollow cylindrical
columns, placed nearly at right angles to the grinding surface, which is
pitted with minute depressions. These teeth are never shed, but are
persistent, and grow on through life, as in the Rodentia, exhibiting in
this respect a striking contrast with those of the Sharks, which are
feeble and numerous, and constantly replaced by rows of successional
teeth.
Fossil teeth of several species, some much larger than the recent,
have been found in the Tertiary, Cretaceous, and Oolitic deposits. The
first British specimen was discovered in the Chalk-marl at Hamsey, in
1820, by myself; but its nature was not suspected until more perfect
examples were obtained from the Kimmeridge Clay at Shotover by Sir
P. Egerton, and were submitted to Dr. Buckland, who subsequently
ascertained their characters and relations by an examination of the
dental organs of a recent Chimæra in the Museum at Leyden in 1835.
[523]

[523] Proceedings of the Geological Society of London, vol. ii. p.

209.
 Lign. 190. Mandible of Edaphodon
Mantelli.
Chalk. Lewes. (1/2 nat. size.)

Lign. 191. Edaphodon leptognathus. 1/2 nat.

London Clay. Bracklesham.
The Upper and Lower Mandibles of the left side; viewed mesially or from
within.

Fig. 1.— The Upper Jaw; left ramus.

p m.—The premaxillary bone.
1, 2, 3.—The three dental tubercles.
2.— The Lower Jaw.
s.—The symphysial surface, by which this ramus is united
to the opposite or right side of the jaw.
4, 5, 6.—The three dental tubercles, or triturating surfaces,
of the left ramus of the lower jaw.

Many specimens, both of the upper and lower mandibles, have

since been discovered in the Eocene beds, Chalk, Upper Greensand,
Galt, Kimmeridge Clay, and Stonesfield Slate. The subject has been
carefully investigated by Sir P. Egerton; and this eminent Ichthyologist
has tabulated the principal forms, and arranged them under five
genera.[524]
[524] Viz. Ganodus, Ischyodus, Edaphodon, Elasmodus, and
Psaliodus. See Quart. Journ. Geol. Soc. vol. iii. p. 35; and Dixon’s
Foss. Sussex.

In some species the external vertical wall of the plate is formed of

hard dentine, resembling enamel; in others the dentine is disposed in
isolated ramifications, producing a dendritical appearance; the
modifications of this structure occasion the differences observable in
the dental plates of the various species. In some, compact dentine
with parallel canals constitutes the mass of the tooth; in others, the
squamous dentine with ramifying tubes prevails.
I have figured the right upper and lower mandibles of the type
named Edaphodon (pavement-tooth), in which there are three teeth or
dental tubercles on each ramus of both jaws, Lign. 191: the lower
mandible is produced anteriorly into a falciform beak:[525] the
articulating surface of the symphysis (2 s) is broad at the base, and
contracts gradually forward until the margins meet at the apex. In
Ischyodus the lower jaw is deeper, less produced in front, and the
margins of the symphysis are parallel until abruptly truncated at the
extremities: the upper jaw has four tubercles on each side.
[525] Hence M. Agassiz proposed the name of Psittacodon
(parrot-tooth) for this genus of Chimæroids.

The upper jaw in Elasmodus has but three tubercles, as in

Edaphodon, but the dentine of which they are composed is confluent,
being rolled round like a scroll in the substance of the bone, one edge
forming the margin of the tooth, and the other being buried deep in its
centre.
The dorsal fin-ray or spine of the Edaphodon is laterally
compressed, with the posterior margin grooved, and the edges armed
with fine teeth: I have a specimen of the spine, with a pair of inferior
mandibles of the same individual, imbedded in a block of chalk from
Kent; by favour of Mrs. Smith, of Tunbridge Wells.
 Lign. 192. Fossil Teeth of Sharks.
Fig. 1.— Hybodus medius. Lias. Lyme Regis.
2.— Hybodus raricostatus. Lias. Bristol.
3.— Carcharodon productus. Tert. Malta.
4.— Hemipristis serra (fragment). Chalk. Ratisbon.
5.— Otodus Obliquus. Tert. Sheppey.

Hybodus. Lign. 192, figs. 1, 2. (Bd. pl. xxviid.)—Intermediate

between the obtuse crushing teeth of the Sharks previously described,
and those sharp, angular, pointed, dental organs of the Squaloids, are
those of the fishes which M. Agassiz has arranged in a sub-family or
group termed Hybodonts; the teeth of which are characterised by their
transversely elongated form, and the series of subacute, compressed,
conical cusps or points, which compose the crown. The median cone is
the principal, the lateral points being shorter and smaller, as in Lign.
192, fig. 2; in some species the difference between the median and
lateral cones is greater, in others less, as in fig. 1. These cusps have a
coating of dense enamel, which is plicated longitudinally on both faces.
The base, which almost equals the crown in size, is composed of a
coarse osseous substance. The internal structure of the crown differs
from that of the Cestracionts, in having no principal pulp-cavity, and in
being chiefly composed of dendritical dentine, with reticulated
medullary canals. The form and organization of these teeth show them
to have been instruments for cutting and tearing food. The Hybodonts,
as we have already stated (p. 581), possessed two spinous dorsal fins;
in their habits and economy they probably did not differ from the
ordinary Sharks. Teeth and spines of this genus are common in the
Trias, Lias, Oolite, and Walden, and occur in the Green Sand and
Chalk. There are several species of teeth and fins in the strata of
Tilgate Forest (Foss. Til. For. pl. x.). In general the teeth are found
detached, but occasionally they occur in their natural position,
adhering to the mineralized cartilaginous jaws (Petrif. Lign. 97); as in
the beautiful fossil figured Bd. pl. xxvii.d; and in the fine specimen of
H. basanus, from the Isle of Wight, figured in the Geol. Soc. Journ. vol.
pl. iv. There are several related genera, founded on the situation, form,
and division of the principal cusps of the teeth.
Sharks with Cutting Teeth.—The jaws of the common squaloid fishes,
as the Lamna (Porbeagle) and Carcharias (Great White Shark), are so
common in collections of natural history, as to render a description
unnecessary. The numerous vertical rows of angular, laterally
compressed, pointed teeth, with sharp or serrated edges—in some
species consisting of a simple trenchant cusp, in others with small
lateral teeth, or denticles, at the base, are characters with which all
are familiar. Fossil teeth of this form are extremely abundant in the
Tertiary and Cretaceous deposits; and are commonly in a beautiful
state of preservation. The genera of these fossil teeth are founded on
the solidity or hollow structure of the cusps, their possessing cutting or
serrated edges, and the presence or absence of lateral denticles. But
the last character cannot in every instance be relied upon, for some
recent Sharks have rows of teeth both with and without denticles.
Carcharodon productus. Lign. 192, fig 3.—The genus Carcharias
comprises the large Sharks with cutting triangular teeth, crenated
(notched) on their margins, and having a broad base. In Carcharodon,
the teeth differ from those of Carcharias in being solid in the centre,
while in the latter they are hollow; but in both genera the teeth exhibit
the same reticulated structure of medullary and calcigerous tubes. The
White Shark and other large species belong to these genera; some of
which are upwards of forty feet in length. But even these colossal
fishes must have been far surpassed in magnitude by the extinct
species of the Tertiary deposits, if the teeth afford a scale of
proportions; for some of the fossil teeth from Malta and the United
States are six inches long, and five wide at the base;[526] being twice
the size of the teeth in the largest living species. The specimen figured
in illustration, Lign. 192, fig. 3, is of a small size.
[526] For instance, some of the Maryland specimens of
Carcharodon megalodon. See an admirable memoir on the Fossil
Squalidæ of the United States, by Dr. R. W. Gibbes, Journ. Acad.
Nat. Science, Philadelphia. At the meeting of the British Association
in 1851, J. S. Bowerbank, Esq. F.R.S. read some interesting
observations on the comparison of these large fossil fishes with the
recent Carcharias glaucus of Australia. See also Owen, Cyclop. Anat.
Art. Teeth.

Hemipristis serra. Lign. 192, fig 4.—The fossil teeth of this genus are
distinguished by serrated edges, that do not extend to the summit,
which is a sharp angular point; as in the fossil represented.

Lign. 193. Fossil Teeth of Sharks. Chalk. Lewes.

Fig. 1.— Corax pristodontus.
2.— Lamna crassidens.
3.— Notidanus microdon.
4.— Ptychodus polygurus; seen laterally.
5.— Ptychodus polygurus; viewed from above.
 6.— Lamna elegans.

Lamna elegans. Lign. 193, fig. 6.—The fishes of the genus Lamna (to
which the recent shark called the Porbeagle belongs) have teeth with
smooth trenchant edges, and a small sharp denticle (little tooth) on
each side the base, as in the fossil, Lign. 193, fig. 6. The specimen,
fig. 2, although devoid of denticles, probably belongs to the same
genus, for reasons already explained. Several species abound in the
Chalk; and they are associated with teeth, which are relatively wider
and shorter, and have large compressed denticles; the latter are
arranged in a separate genus, named Otodus (eared-tooth), by M.
Agassiz. The specimen figured Lign. 192, fig. 5, represents O.
obliquus; another species, Otodus appendiculatus, is abundant in the
Sussex Chalk. The large, wide, triangular, smooth teeth, with trenchant
edges, and destitute of lateral denticles, so common in the Chalk, are
related to Lamna, and are comprised in the genus Oxyrhina (Poiss.
Foss. tom. v. tab. xxxiii.).
Notidanus microdon. Lign. 193, fig. 3.—These teeth differ remarkably
from those of the other genera of Sharks. The crown of each tooth is
composed of a series of sharp angular enamelled points, the first of
which is the largest, and is notched on its anterior edge; the base or
root is osseous, flat, with a slight longitudinal depression below the
border of enamel. These teeth are comparatively rare in the Chalk.
One species has been found in the Oxford Clay; and several in the
Tertiary strata. Specimens occur in Hordwell Cliff.
Corax pristodontus. Lign. 193, fig. 1.—The teeth of the fossil Corax
chiefly differ from those of the recent genus Galeus, to which the
Tope, or Grey Shark, belongs, in being solid; they are of a triangular
form, with a deep concavity or notch on the posterior margin, the base
of which is prolonged and forms three or four angular points: the
anterior edge of the tooth is finely serrated. The root of the tooth, as
in Notidanus, is a broad osseous plate. There is much diversity of form
in the Chalk specimens, which are all of a small size, as in Lign. 193,
fig. 1. In Sussex they are more common in the Chalk-marl than in the
Chalk.
 The only fossil teeth of the Shark family resembling those of the
tertiary Carchariodonts, that have been discovered in the strata below
the Chalk, are from the carboniferous deposits of Yorkshire and
Armagh. These teeth are compressed, triangular, crenated on the
edges, with large plaits or folds on the enamelled surface, towards the
base of the crown. M. Agassiz refers them to a new genus, viz.
Carcharopsis, with the specific name of Prototypus.
Fossil Vertebræ of Sharks.—As the cartilaginous nature of the
skeleton in this family renders it unfavourable to preservation in the
mineral kingdom, the durable parts already described, and those which
are ossified, are almost the only relics found in a fossil state. The
dermal integument is, however, sometimes preserved; and I had a
beautiful example of shagreen, composed of irregular minute
hexagonal scales, one of which is represented highly magnified, Lign.
185, fig. 1.
In the Galeus and Carcharias the vertebræ are more ossified than
in many other genera of cartilaginous fishes, and fossil vertebræ of
these sharks often occur in the cretaceous and other strata. Groups of
vertebræ of a large size occasionally occur in the Sussex Chalk; they
are circular, biconcave, and very short; one specimen is four inches in
diameter, and one inch long; their concavities are consequently
shallow. These vertebra: are composed of two shallow conical disks,
which are united by their summits, at the axis, and are connected and
supported by numerous wedge-shaped plates, that radiate from the
centre to the periphery (see Foss. South D. pl. xxxiii. fig. 10). My
collection contained a connected series of forty small vertebræ from
the Chalk near Lewes, which probably belonged to the same species of
Shark as the dorsal spine named Spinax major (Poiss. Foss. tom. iii. pl.
xla fig. 6).
Squaloraia. In the Lias of Lyme Regis, that inexhaustible storehouse
of fossil treasures, a considerable portion of the skeleton of a very
remarkable fish, partaking of the characters of the Sharks and Rays,
was discovered by Miss Mary Aiming, and is now in the Museum of the
Bristol Institution.[527] In this fish the jaws are prolonged into a beak,
like the Pristis (Saw-fish). It has the head of a Shark, with a long beak;
vertebræ of the Rays; with pectoral and ventral fins, almost equally
developed; a tail armed with, a spine; and spinous bosses, as in the
true Rays.
[527] It is figured and described by Dr. Riley, Geol. Trans. 2d ser.
vol. v. pl. iv. See also Poiss. Foss. tom. iii. pl. xlii.

Fossil Pristis, or Saw-fish.—This well-known predatory fish, which is

allied to the Rays and Sharks, and referred by M. Agassiz to the family
of Raiidæ, has projecting from its snout an osseous, flat, horizontal
plate, or beak, equal in length to one-third of the fish, and armed on
each side by a row of elongated, compressed, pointed teeth, implanted
in sockets; the front margin of these teeth is convex, the posterior
concave; this defence is termed the saw, and constitutes a most
powerful weapon. The Pristis has also numerous small obtuse teeth on
the jaws. The remains of the beak, or saw, of an extinct species of
Pristis have been discovered in the Bagshot Sand at Goldsworth Hill,
Surrey,[528] and three other species have been found in the London
clay of the Isle of Sheppey, and the Eocene beds at Bracklesham and
Hordwell.[529]
[528] Proc. Geol. Soc. vol. ii. p. 687.
[529] Two teeth are figured in Dixon’s Fossils of Sussex, pl. xii.;
the specimens are in the British Museum; see Petrifactions, p. 414.

Fossil Rays.—The teeth of these fishes are characterised by the

extraordinary transversal development of the median teeth in both
jaws. Instead of pointed teeth, they have wide, flat, tesselated dentary
plates in each jaw, composed of distinct pieces, juxtaposed and
connected by their margins, and united by fine sutures. In some
species the teeth are equal, in others of various sizes; they present
numerous modifications of arrangement, and are always disposed in
symmetrical rows. In the genus Myliobatis (Eagle-ray) the teeth of the
median row are of an extraordinary width, while their length does not
exceed that of the lateral plates, or chevrons, which are of an irregular
hexagonal form, and disposed in two or three rows on each side.
There are five living species of Myliobatis, and eighteen fossil species
have been found in the Tertiary strata at the Isle of Sheppey, Hordwell
Cliff, and Bracklesham Bay. I have figured a specimen of part of the
upper jaw of a species (M. micropleurus, Lign. 194, fig. 2), in which
the median teeth are very wide, and have two lateral rows of small,
irregularly hexagonal, plates. The surface of the teeth in this species is
smooth; but in others it is striated longitudinally (Bd. pl. xxvid. fig. 14).
In an allied genus, Ætobatis, from the Tertiary beds at Bracklesham,
the lower jaw projects beyond the upper, and in each there is a row of
flat, transverse teeth, without lateral plates.

Lign. 194. Fossil Teeth of Fishes

Fig. 1. Ceratodus emarginatus: 1/2 nat. Lias Bone-
— bed. Aust Cliff. A single dental plate.
2. Myliobatis micropleurus: 1/3 nat. Tert. Isle
— of Sheppey. A series of six median teeth,
with the corresponding lateral teeth.
3. Pycnodus Mantelli. Weald. Tilgate Forest.
— This specimen consists of the vomerine
bone, with a median row of flat, arched
teeth, and two rows on each side of
elliptical teeth, arranged alternately.

To this notice of the fossil Rays, we may add, that a gigantic

Torpedo has been discovered in that celebrated locality of Ichthyolites,
Monte Bolca: and that Sir Philip Egerton has recently enriched his
matchless collection of fossil fishes, by a perfect Ray, from Mount
Lebanon. It is figured and described in the Quarterly Geol. Journ. vol.
i. pl. v. p. 225, under the name of Cyclobatis oligodactylus. It is a small
species, resembling the common Rays in its general appearance, but is
surrounded by a broad, flexible, cartilagino-membranous fin; the skin
is smooth, the teeth and eyes are small, the tail is slender; there are
no traces of dermal spines, tubercles, or defensive weapons. In many
points of structure it resembles the Torpedo; and may possibly, like
that fish, have possessed an electric organ. It is 31/2 inches long, and 3
inches wide across the expanded fins.
 CHAPTER XV.
FOSSIL ICHTHYOLOGY; COMPRISING THE
GANOID, CTENOID, AND CYCLOID FISHES.
The fishes we have hitherto examined belong to the first order, the
Placoidians; we now pass to the fossil remains of the second order, the
Ganoidians, which are distinguished by their brilliant angular scales,
formed of osseous or horny plates, densely covered with enamel. This
order contains six or more families, comprising many genera and
numerous species; our investigation must be restricted to a selection
from the principal genera of the Ganoids, properly so called, and of the
Sauroids, or lizard-like fishes.[530]
[530] The fishes of these orders are described in Poiss. Foss.
tom. ii.

The first family, termed Lepidoides, contains several genera, which

are defined as possessing either numerous rows of brush-teeth, or of
obtuse conical teeth; flat, rhomboidal scales, arranged parallel with the
body; and an osseous, or partially osseous, skeleton. In one division of
this family, the body is either elongated or fusiform, the mouth
furnished with brash-teeth only, and the tail heterocercal, or unequally
bilobed (see p. 576). To this group belong several genera, which are
restricted to the Secondary formations more ancient than the Oolite;
while the other group, with homocercal tails, lived in the Oolitic and
Cretaceous seas. Two genera, in particular, abound in the Permian and
Carboniferous strata; namely, Amblypterus and Palæoniscus.[531]
[531] For the characters, affinities, and distribution of these and
the allied genera of the Heterocerque Ganoid fishes, see Sir P.
Egerton’s Memoir in the sixth volume of the Quarterly Geological
Journal.
 Amblypterus. Lign. 187. Wond. p. 740. Bd. pl. xxvii.b.—The fishes of
this genus, as the name indicates, have very large and wide fins,
composed of numerous rays. The scales are rhomboidal and finely
enamelled; the tail is heterocercal. The figures referred to convey a
correct idea of the form and external characters. Beautiful pyritous
imprints of Amblypteri occur in the Carboniferous slate of Saarbrück, in
Lorraine; and fine specimens in the ironstone nodules of the same
locality. On the shore at Newhaven, near Leith, similar fossils occur in
nodules washed out of the cliffs of coal-shale (Bd. p. 278).
Palæoniscus. Ly. p. 304.—The fishes of this genus differ from those
of Amblypterus in the relatively moderate size of the scales, and the
numerous little rays on their margins. They have rhomboidal scales,
which in some species are very small, and in others large. They have
numerous brush-teeth. Several peculiar species, found in the marl-
slates and magnesian limestones of the Permian system, are very
widely distributed, occurring in the British Isles, Germany, and the
United States.
In some localities the small species occur in groups; on a slab of
red sandstone, in the Museum of the Geological Society, from Tyrone,
between two and three hundred perfect fishes (P. catopterus) are
imbedded on a space not exceeding two feet square.
A remarkable circumstance relating to the fishes of this genus is
the almost constant absence of the bodies of the vertebræ in
otherwise well-preserved specimens, and in which the spinal processes
and the ribs are entire: occasionally, however, examples occur with
some of the vertebræ perfect. An explanation of the above
phenomenon may perhaps be found in the probable originally
cartilaginous nature of the bodies of the vertebræ, and the osseous
structure of the enduring apophyses and ribs;[532] while those rare
specimens which possess a few bony vertebræ may be regarded as
exceptions, in which ossification took place in a structure essentially
cartilaginous.
[532] Professor Owen states that a similar condition of the spinal
column obtains in the fossil Microdonts.—Rep. Brit. Assoc. 1846.
 The fishes found in the copper-schists of the Zechstein, at Mansfeld
in Saxony, are generally impregnated with copper pyrites, and their
scales are as brilliant as burnished gold. These ichthyolites are almost
always in contorted and twisted positions; which appearance M.
Agassiz attributes to contraction of the muscular tissues after death,
during the progress of decomposition, and before the fishes sank down
and became imbedded in the mud. (Poiss. Foss. tom. ii. p. 70.)
The fishes of the genus Palæoniscus are often found in the shales
and marls of the Permian and Carboniferous systems of England and
Scotland. At East Thickley, in the county of Durham, numerous
specimens have been found.[533] The lower Carboniferous strata at
Burdie-house, a locality we have before mentioned, have yielded
several species of Palæoniscus, associated with teeth and other
remains of large sauroid fishes.[534] On the continent also they prevail
in deposits of the same epoch; Eisleben and Mansfeld, iii Saxony, are
well-known localities. In North America they have been discovered in
strata of probably the same age.[535] In fine, the genera Amblypterus
and Palæoniscus may be regarded as characteristic "medals" of the
geological epoch which intervened between the Devonian and Triassic
formations.
[533] See Professor Sedgwick on the Magnesian Limestone.
Geol. Trans. 2d ser. vol. iii.; and Prof. King’s Monograph on the
Permian Fossils, published by the Palæontographical Society.
[534] Dr. Hibbert’s Memoir on the Fossils of Burdie-house.
[535] Geology of Massachusetts, by Professor E. Hitchcock.

We will next examine a few genera of the homocercal Lepidoids

and Pycnodonts, whose relics are chiefly distributed in the Lias, Oolite,
Purbeck, and Wealden.
 Lign. 195. Outline of the Dapedius. Lias. (1/2 nat. size).

Dapedius. Lign. 195.—At Lyme Regis, and other productive localities

of the fossils of the Lias, large masses of angular enamelled scales,
and occasionally entire specimens of the fishes to which they
belonged, have for many years been collected. Sir H. De la Beche first
scientifically investigated the structure of these Ichthyolites, and
pointed out their characters and relations. The Dapedius (of which a
restored figure is given in Lign. 195) is a flat, laterally compressed fish,
with a rounded head, and fins of moderate size. The body rapidly
contracts towards the root of the tail, the fin of which is large and
symmetrically lobed. The mouth is furnished with several rows of small
conical teeth, which are crenated at their summits, and has brush-
teeth on the palatine bones; the jaws are short. The scales are
rhomboidal, highly polished, and united laterally by short processes; as
in many other ganoid fishes. It belongs to the Lepidoids.
The Tetragonolepis is a Liassic fish, very similar in shape to the
Dapedius. It was formerly grouped with the Lepidoidei, but Sir P.
Egerton has lately discovered that it essentially differs from those
fishes in the character of its scales and teeth, and that it belongs to
the Pycnodonts.

Lepidotus.[536] Lign. 186, 196, 197.—Scales of a dark-brown, almost

black colour, with a glossy enamelled surface, and of a rhomboidal or
lozenge form, and teeth equally dark and glossy, of an obtuse
hemispherical figure, are very common in the Wealden strata of the
south-east of England and in the Isle of Purbeck. They are called by
the quarry-men fishes' scales and eyes. The collectors of the last
century used to term the obtuse circular teeth of this and the related
genera Bufonites, from a supposition that they were formed in the
heads of toads. These relics belong to an extinct genus named
Lepidotus, which contains numerous species, that are distributed in the
Oolite, Purbeck, and Wealden formations. These fishes resembled the
Carps in their general form, but they have no anatomical relations to
that family. The body is covered with large rhomboidal scales, which
are protected on the external surface by a thick plate of enamel (Lign
196, fig. 3). The lateral line, which is slightly arched, passes direct
from the operculum to the middle of the insertion of the caudal fin.
The head, and even the face, are cased with osseous and enamelled
plates. The bones of the surface of the skull are very large, and are
connected by sinuous sutures. The jaws are short and rounded, and
furnished with a row of obtuse, conical, circular teeth (see Lign. 197),
and several rows of sessile teeth, more or less contracted at the base,
which forms a very short pedicle that is anchylosed to the bone. The
fossil Lepidoti are found, for the most part, in fluviatile deposits, as in
the Purbeck and Wealden strata; and it is probable they inhabited the
rivers or sea-coasts, and not deep waters.
[536] Poiss. Foss. tom. ii. p. 233.
 Lign. 196. Scales Fin of Lepidotus Mantelli.
and
Wealden.
Tilgate Forest. (Nat. size.)
Fig. 1.
Scale, with a single process of attachment.
—
2.
One of the scales of the dorsal line.
—
3. Scale (external surface), with a bifurcating
— process of attachment; the enamelled portion
(which alone is visible when the scales are in
their natural position on the animal) has
longitudinal grooves or folds.
4. Scale (viewed on the inner surface), having a
— bifurcating process of attachment and a tooth
or projection on each side, to connect the scale
laterally with the adjoining scales.
5. The front ray of the dorsal fin, covered with two
— rows of enamelled scales, and another ray
behind it.

The scales and teeth figured Lign. 196, 197, belong to the larger
species of the Wealden. The remains of this fish were first collected in
Tilgate Forest, and several teeth and: scales are figured Foss. Til. For.
pl. v. and x.; considerable portions of connected scales have since
been found; also the head entire, and the fins more or less perfect. A
specimen in my collection retained a mass of the scales near the
insertion of the tail, a foot wide; indicating the original to have been
twelve feet long, and its body three feet broad. The scales are
distinguished from other species by the folds or grooves on their
enamelled surface; and the teeth by the contracted base, or pedicle,
which is a little narrower than the crown (Lign. 197, and Pl. VI. fig.
10). A species (L. Fittoni) closely related to the above is equally
abundant in the Weald of Sussex; the scales are not striated, and the
teeth have no pedicle.

Lign. 197. Portion of the Jaw of Lepidotus.

Wealden. Tilgate Forest. (Nat. size.)
This specimen shows three successional teeth
beneath a row of teeth in use.

The intimate structure of the teeth of the Lepidotus is beautifully

preserved, and may be easily examined in thin transverse and vertical
sections, viewed by transmitted light: see Pl. VI. fig. 10. The dentine is
composed of bundles of tubes, continued from the cells of the osseous
base, radiating in a vertical direction to the surface of the tooth, as
seen in Pl. VI. fig. 10, and giving off branches at an acute angle; but
when more highly magnified, the finer branches are seen to be spread
out and arched at their extremities, "presenting the appearance of the
stems of corn, beaten down by heavy rain."[537]
[537] Odontography, p. 70. See the beautiful representation of
this structure, pl. xxxi.

The dorsal and pectoral fins of these fishes are very strong, and
consist of several bony rays. There is a double row of acuminated
enamelled scales, arranged more or less obliquely, on the anterior
margin of the dorsal and anal fins, and on both margins of the caudal:
part of the first ray of a dorsal fin, with scales, is represented Lign.
196, fig. 5.
A small species of Lepidotus (A. minor) is common in the Purbeck
limestone, and specimens may often be procured from the quarries
near Swanage;[538] it has also been found at Hildesheim, in Saxony, by
M. Roemer. The detached scales abound in the limestones; and the
splendid fossil reptile from Swanage, figured Petrif. Lign. 38, is
sprinkled with the scales and minute teeth of this fish.
[538] Fine specimens of this fish are in the British Museum
(Petrifactions, p. 429), and in the Museum at Dorchester.

The majority of the species of Lepidotus belong to the Lower

Oolites and the Lias. The habits of the Lepidoti, as indicated by the
form and structure of the teeth, were those of fishes whose food
consisted of crustaceans, shelly mollusca, &c.; for the dental organs
are peculiarly adapted for the crushing and grinding of such
substances; and the teeth of the adult fishes are generally worn down
by use.
Pycnodus. Pl. I. fig. 3; Lign. 194, fig. 3.—The fishes of the family of
Pycnodonts, so named from the thickness of their teeth, have an
osseous skeleton, a flat body covered with rhomboidal scales, and flat
or rounded teeth disposed in several rows on the palatine, vomerine,
intermaxillary, and premandibular bones.[539] As in the Lepidotus,
these teeth are constructed for crushing, and have generally a smooth,
dense, convex or flattened crown, with a highly polished surface. This
genus belongs chiefly to the Oolite; it is found also in the Lias, Chalk,
and Wealden. A perfect fish of the genus Pycnodus (P. rhombus), from
the Jura limestone, at Torre d’Orlando, near Castellamare, is figured in
the frontispiece of the first volume of this work; and a vomerine bone,
with teeth, from Tilgate Forest, in Lign. 194, fig. 3. In the last fossil
there is a median row of flat,, elongated, transversely arched, smooth,
glossy teeth, with a double alternate row of small sub-circular teeth on
each side, attached to the bone, which is imbedded in Tilgate grit.
Specimens of this kind, belonging to one or more species of Pycnodus,
occur in the Wealden of Sussex; they were among my earliest
discoveries in Tilgate Forest (Foss. Til. For. pl. xvii. figs. 26, 27).
Examples are met; with in which all the teeth are shed, and the bony
plate of the vomer alone remains.
[539] The intermaxillary, palatine, and vomerine bones compose
the vault or roof of the mouth; the vomer occupying the middle; the
intermaxillary the front; and the palatine bones the sides; the
premandibular bones belong to the lower border of the mouth.

Gyrodus. Lign. 198.—In another genus of the Pycnodonts, termed

Gyrodus, the crowns of the teeth are deeply furrowed, the structure of
the dentine is very dense, and the pulp-cavity large and simple. One
species occurs in the Speeton clay of Yorkshire, and another in the
Sussex weald; but the teeth are chiefly found in the Oolite and Chalk.
As in Pycnodus, the teeth are distributed in rows on the bones
composing the roof, floor, and sides of the mouth.[540]
[540] An extraordinarily perfect lower jaw of a Gyrodus is to be
seen in the British Museum (Foss. Brit. Mus. p. 439).

These characters are beautifully displayed in the Russian specimen,

Lign. 198. This interesting fossil was presented to me by the late
Stephen Cattley, jun. Esq., who collected it in 1839, in a valley near
Rjeff, a village on the banks of the Volga. Mr. Cattley informed me
"that many fossils are found in that and the neighbouring valleys; and
the locality is frequented by Russian geologists when the season
permits, which is but seldom, owing to the long duration of the snow,
and the heavy rains which accompany the thaw." This specimen
consists of the vomerine bone, which is of a coarse texture, and five
rows of teeth; the median row consists of very large elliptical teeth;
those of the lateral rows are much smaller and arranged alternately.
The peculiar structure of the teeth of this genus[541] is finely displayed
in this fossil. The ample, deep, and simple pulp-cavity is seen in
several teeth, where the crown of dentine has been worn off, filled
with a pure white calcareous spar; one of these cavities is marked a.
The dentine is extremely dense, consisting of very minute calcigerous
tubes, and passes into an external layer of enamel.
 [541] Odontography, p. 72.

Lign. 198. Gyrodus Murchisoni[542] (G. A. M.)

Oolite? Russia.
(Collected by the late Stephen Cattley, jun.
Esq.)
Fig. 1. The vomerine bone of a fish, with five
— rows of teeth; seen from above; many
of the crowns of the teeth are worn
away by use, and the large pulp-
cavities, filled with white spar, are
exposed; as at a.
2. Lateral view of the same.
— a. One of the pulp-cavities filled with
spar.

[542] It is with peculiar pleasure that I inscribe this new species

of Gyrodus to Sir R. I. Murchison, in commemoration of those
extended and successful geological researches in the Russian
empire, which have conferred additional honour on his distinguished
name.

The fishes of the genus Gyrodus have the body large, flat, and
elevated; the dorsal and anal fins are very long; and the tail is forked,
with equal elongated lobes. The scales are laterally connected by
strong processes, as in Lepidotus.
 Other genera related to the foregoing occur in the Oolite; as for
example, Microdon, thus named from the smallness of its very
numerous flat angular teeth, arranged in many rows; Placodus, in
which the teeth are few, flat, and of great size;[543] and Platysomus
(flat-body), with orbicular, clavate, teeth.
[543] Odontography, pl. xliii. fig. 1, and pl. xxx. fig. 2.

In these fishes, also, the dental organs are well adapted for the
comminution of shell-fish, and other hard bodies.
Cephalaspides of the Devonian System.—The remains of the three
genera of Ganoid fishes that we have now to notice are of a very
remarkable character, and are found exclusively in the Devonian or Old
Red system; most frequently in Scotland, but also in other parts of the
British Isles, and in Europe and America. These fishes agree in one
general character, that of having extensive osseous plates, or
scutcheons; their general aspect will be understood by reference to
Lign. 199, 200, 201. There are no vestiges of the bodies of the
vertebræ, which, therefore, were probably cartilaginous. These fishes
constitute a distinct family with the name Cephalaspides, from the
character of the first genus we propose to describe.

Lign. 199. Cephalaspis Lyellii. (1/4 nat. size.)

Devonian. Forfarshire.
Crushed specimen; seen from above.
 Lign. 200. Cephalaspis Lyellii. (1/4 nat. size.)
Devonian. Glammis, Forfarshire.
Lateral view, showing the produced dorsal lobe
of the tail.

Cephalaspis Lyellii. Lign. 199, 200.—The most striking feature in the

Ichthyolites of this genus is the enormous scutcheon, or buckler, which
forms the head, and is prolonged posteriorly into two lateral horns or
points; this part so closely resembles the cephalic shield of certain
trilobites (see Lign. 175), that the first found specimens were
supposed to be the remains of unknown crustaceans. The name
Cephalaspis (buckler-head) is derived from this character. This
remarkable appearance is occasioned by the intimate anchylosis of all
the bones of the cranium. The body of these fishes is relatively smaller
than the head; it has one dorsal fin, and terminates in a tapering tail,
supporting a fin. There are two small eyes, placed towards the middle
of the head. The body is covered with rhomboidal scales; and the head
with discoidal scales, which are highly ornamented with radiated
markings[544] (Ly. p. 344, fig. 396). There are four species of
Cephalaspis at present known.
[544] Poiss. Foss. tom. ii. p. 135.

The other genera are equally unlike any recent types of the class of
fishes. No perfect examples have been found, and some parts of their
structure are still unknown; the annexed figures, Lign. 201, have been
drawn by Mr. Dinkel (the eminent artist employed by M. Agassiz), with
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