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Hazzard's
GERIATRIC MEDICINE
and GERONTOLOGY
EIGHTH EDITION
Jeffrey B. Halter • Joseph G. Ouslander • Stephanie Studenski

Me Kevin P. High • Sanjay Asthana • Mark A. Supiano
Graw
Hill Christine S. Ritchie • Kenneth Schmader
Notice
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Eighth Edition
Hazzards
Geriatric Medicine
and Gerontology
Editors Mark A. SupianorMD
I). Keith Barnes, MD and Dottic Barnes Presidential
Jeffrey B. Halter, MD Endowed (.'hair in Medicine
Professor Emeritus Professor and Chief, Division of Geriatrics
Deportment of Internal Medicine University of Utah School of Medicine
Division ofGeriatric andPalliativeMedkine Executive Director, University of Utah Center on Aging
University of Michigan Salt lake City, Utah
Parkway Visiting Professor in Geriatrics
Department of Medicine .
Christine S. Ritchie, MD MSPH
Yong Loo Lin School of Medicine .
Kenneth L Minaker Endowed Chair in Geriatric Medicine
National University of Singapore .
Director Mongan Institute Center for Aging and Serious Illness
Ann Arbor, Michigan Professor of Medicine
Massachusetts General Hospital and
Joseph G. Ouslander, MD Harvard Medical School
Professor of Geriatric Medicine Boston, Massachusetts
Senior Advisor to the Dean tor Geriatrics
.
Charles E Schmidt College of Medicine Kenneth Schmader, MD
.
Professor ( Courtesy ), Christine E Lynn College of Nursing Professor of Medicine-Geriatrics
Florida Atlantic University Co- Director, Pepper Older Americans Independence ('enter
Boca Raton, Florida Duke University Medical Center
Editor- in-Chief, Journal of the American Geriatrics Society .
Director Geriatric Research Education and Clinical Center
Durham YA Health Care System
Stephanie Studenski,MD, MPH
Professor Emeritus
.
Durham North Carolina
University of Pittsburgh School of Medicine Editor Emeritus and Senior Advisor

Pittsburgh, Pennsylvania
William R. Hazzard, MD
Kevin P. High, MD,MS Professor Emeritus, Internal Medicine — Gerontology and
President, Atrium Health — Wake Forest Baptist Geriatric Medicute
Professor of Internal Medicine-Infectious Diseases I. Paul Sticht Center on Aging and Rehabilitation
Wake Forest School of Medicine Wake Forest School of Medicine
Winston- Salem, North Carolina Winston‘Salem,North Carolina
Sanjay Asthana, MD, FACP Senior Editorial Assistant
Associate Dean lor Gerontology
Professor and Head, Division of Geriatrics and Gerontology
Nancy F.Woolard
.
Director N1A/NIH Wisconsin Alzheimer Disease Senior Manager tor Clinical Research
Section on Gerontology & Geriatric Medicine
Research Center ( ADRC)
.
Director Madison VAMC Geriatric Research . Wake Forest School of Medicine
Winston - Salem, North Carolina
Hducation and Clinical Center (GRECC)
Duncan G ami Lottie II, BalLintine Endowed Chair in Geriatrics
University of Wisconsin School of Medicine and Public Health
.
Madison Wisconsin
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Contents
Contributors
Foreword
Preface
Part I
Principles of Gerontology
1 Biology of Aging and Longevity
David B. Lombard, Richard A. Miller, Scott D. Pletcher
2 Demography and Epidemiology
Michelle C. Odden, Kendra D. Sims, Anne B. Newman
3 Immunology and Inflammation
Albert C. Shaw, Thilinie D. Bandaranayake
4 Psychosocial Aspects of Aging
Steven M. Albert, Cynthia Felix
5 Sex Differences in Health and Longevity
Steven N. Austad
6 Social Determinants of Health, Health Disparities, and Health Equity
Laura Block, W. Ryan Powell, Andrea Gilmore-Bykovskyi, Amy J. H. Kind
Part II
Principles of Geriatrics
SECTION A: Assessment
7 Decision Making and Advance Care Planning: What Matters Most
Daniel D. Matlock, Hillary D. Lum
8 Principles of Geriatric Assessment
David B. Reuben, Ryan J. Uyan, Valerie S. Wong
9 Mental Status and Neurologic Examination
James E. Galvin, Michelle M. Marrero
10 Assessment of Decisional Capacity and Competencies
Margaret A. Drickamer, Sarah Stoneking
11 Prevention and Screening
Ashwin A. Kotwal, Sei J. Lee
SECTION B: Age-Friendly Care Across Settings
12 Age-Friendly Care
Terry Fulmer, Maryama Diaw, Chaoli Zhang, Jinghan Zhang, Wendy Huang, Amy Berman,
Tara Asokan, Kedar S. Mate, Leslie Pelton
13 Geriatrics Around the World
Hidenori Arai, Jacqueline C. T. Close, Len Gray, Finbarr C. Martin, Luis Miguel Gutierrez
Robledo, Stephanie Studenski
14 Models of Hospital and Outpatient Care
Jonny Macias Tejada, Michael L. Malone
15 Emergency Department Care
Christopher R. Carpenter, Ula Hwang
16 Institutional Long-Term and Post-Acute Care
Joseph G. Ouslander, Alice F. Bonner
17 Community-Based Long-Term Services and Support, and Home-Based Medical Care
Jessica Colburn, Jennifer Hayashi, Bruce Leff
18 Transitions of Care
Elizabeth N. Chapman, Andrea Gilmore-Bykovskyi, Amy J. H. Kind
19 Value-Based Care
David J. Meyers, Heidi Wold, Joseph G. Ouslander
20 The Role of Social Workers
Ruth E. Dunkle, Jay Kayser, Angela K. Perone
21 The Patient Perspective
Preeti N. Malani, Erica S. Solway, Jeffrey T. Kullgren
SECTION C: Special Management Issues
22 Medication Prescribing and De-Prescribing
Paula A. Rochon, Sudeep S. Gill, Christina Reppas-Rindlisbacher, Nathan M. Stall, Jerry H.
Gurwitz
23 Substance Use and Disorders
Benjamin H. Han, Alexis Kuerbis, Alison A. Moore
24 Integrative Medicine and Health
Julia Loewenthal, Gloria Y. Yeh, Darshan H. Mehta, Peter M. Wayne
25 Patient-Centered Management of Chronic Diseases
Caroline S. Blaum, Aanand D. Naik
26 Legal Issues
Marshall B. Kapp
SECTION D: Surgical Management
27 Perioperative Care: Evaluation and Management
Shelley R. McDonald
28 Anesthesia
Leanne Groban, Chandrika Garner
29 Surgical Quality and Outcomes
Hiroko Kunitake
SECTION E: Nutrition
30 Nutrition Disorders, Obesity, and Enteral/Parenteral Alimentation
Dennis H. Sullivan, Larry E. Johnson, Jeffrey I. Wallace
31 Disorders of Swallowing
Nicole Rogus-Pulia, Steven Barczi, JoAnne Robbins
32 Oral Health
Joseph M. Calabrese, Judith A. Jones
SECTION F: Sensory Function
33 Low Vision: Assessment and Rehabilitation
Gale R. Watson, Katharina V. Echt
34 Hearing Loss: Assessment and Management
Su-Hua Sha, Kara C. Schvartz-Leyzac, Jochen Schacht
SECTION G: Gender and Sexuality
35 Sexuality, Sexual Function, and the Aging Woman
Monica Christmas, Kaitlyn Fruin, Stacy Tessler Lindau
36 Gynecologic Disorders
Thomas Clark Powell, Russell Stanley, Holly E. Richter
37 Sexuality, Sexual Function, and the Aging Man
J. Lisa Tenover, Alvin M. Matsumoto
38 Benign Prostate Disorders
Catherine E. DuBeau, Christopher D. Ortengren
Part III
Geriatric Conditions
39 Systems Physiology of Aging and Selected Disorders of Homeostasis
George A. Kuchel
40 Applied Clinical Geroscience
Sara E. Espinoza, Jamie N. Justice, John C. Newman, Robert J. Pignolo, George A. Kuchel
41 Managing the Care of Patients with Multiple Chronic Conditions
Stephanie Nothelle, Francesca Brancati, Cynthia Boyd
SECTION A: Geriatric Syndromes
42 Frailty
Luigi Ferrucci, Jeremy D. Walston
43 Falls
Stephen R. Lord, Jasmine C. Menant
44 Sleep Disorders
Armand Ryden, Cathy Alessi
45 Syncope and Dizziness
Ria Roberts, Lewis A. Lipsitz
46 Pressure Injuries
Joyce M. Black
47 Incontinence
Camille P. Vaughan, Theodore M. Johnson, II
48 Elder Mistreatment
Mark S. Lachs, Tony Rosen
SECTION B: Mobility
49 Muscle Aging and Sarcopenia
Alfonso J. Cruz-Jentoft
50 Mobility Assessment and Management
Valerie Shuman, Caterina Rosano, Jennifer S. Brach
51 Osteoporosis
Gustavo Duque, Mizhgan Fatima, Jesse Zanker, Bruce R. Troen
52 Osteoarthritis
Michele R. Obert, Ernest R. Vina, Jawad Bilal, C. Kent Kwoh
53 Hip Fractures
Ellen F. Binder, Simon Mears
54 Therapeutic Exercise
Kerry L. Hildreth, Kathleen M. Gavin, Christine M. Swanson, Sarah J. Wherry, Kerrie L.
Moreau
55 Rehabilitation
Cynthia J. Brown
SECTION C: Mentation
56 The Aging Brain
Luigi Puglielli
57 Cognitive Changes in Normal and Pathologic Aging
Bonnie C. Sachs, Brenna Cholerton, Suzanne Craft
58 Delirium
Matthew E. Growdon, Tanya Mailhot, Jane S. Saczynski, Tamara G. Fong, Sharon K.
Inouye
59 Dementia Including Alzheimer Disease
Cynthia M. Carlsson, Nathaniel A. Chin, Carey E. Gleason, Luigi Puglielli, Sanjay Asthana
60 Behavioral Symptoms of Dementia and Psychoactive Drug Therapy
Carol K. Chan, Constantine G. Lyketsos
61 Parkinson Disease and Related Disorders
Vikas Kotagal, Nicolaas I. Bohnen
62 Cerebrovascular Disease
Nirav R. Bhatt, Bernardo Liberato
63 Other Neurodegenerative Disorders
John Best, Howie Rosen, Victor Valcour, Bruce Miller
64 Traumatic Brain Injury and Chronic Traumatic Encephalopathy
Ann C. McKee, Daniel Kirsch
65 Major Depression
Whitney L. Carlson, William Bryson, Stephen Thielke
66 General Topics in Geriatric Psychiatry
Ellen E. Lee, Jeffrey Lam, Dilip V. Jeste
Part IV
Principles of Palliative Medicine and Ethics

67 Palliative Care and Special Management Issues
Paul Tatum, Shannon Devlin, Shaida Talebreza, Jeanette S. Ross, Eric Widera
68 Pain Management
Roxanne Bavarian, Amber K. Brooks
69 Management of Common Nonpain Symptoms
Christine S. Ritchie, Alexander Smith, Christine Miaskowski
70 Palliative Care Across Care Settings
Lisa Cooper, Laura Frain, Nelia Jain
71 Effective Communication Strategies For Patients with Serious Illness
Brook Calton, Matthew L. Russell
72 Ethical Issues
Timothy W. Farrell, Caroline A. Vitale, Christina L. Bell, Elizabeth K. Vig
Part V
Organ Systems and Diseases

SECTION A: Cardiovascular System
73 The Aging Cardiovascular System
Ambarish Pandey, George E. Taffet, Dalane W. Kitzman, Bharathi Upadhya
74 Coronary Heart Disease and Dyslipidemia
Michael G. Nanna, Karen P. Alexander
75 Valvular Heart Disease
Nikola Dobrilovic, Dae Hyun Kim, Niloo M. Edwards
76 Heart Failure
Mathew S. Maurer, Scott L. Hummel, Parag Goyal
77 Cardiac Arrhythmias
Nway Le Ko Ko, Win-Kuang Shen
78 Peripheral Vascular Disease
Jonathan R. Thompson, Jason M. Johanning
79 Hypertension
Mark A. Supiano
SECTION B: Pulmonary
80 Respiratory System and Selected Pulmonary Disorders
Daniel Guidot, Patty J. Lee, Laurie D. Snyder
81 Chronic Obstructive Pulmonary Disease
Carolyn L. Rochester, Kathleen M. Akgün, Jennifer D. Possick, Jennifer M. Kapo, Patty J.
Lee
SECTION C: Nephrology
82 Aging of the Kidney
Jocelyn Wiggins, Abhijit S. Naik, Sanjeevkumar R. Patel
83 Kidney Diseases
Mark Unruh, Nitin Budhwar
SECTION D: Gastroenterology
84 Aging of the Gastrointestinal System and Selected Lower GI Disorders
Karen E. Hall
85 Upper Gastrointestinal Disorders
Alberto Pilotto, Marilisa Franceschi
86 Hepatic, Pancreatic, and Biliary Diseases
Dylan Stanfield, Mark Benson, Michael R. Lucey
87 Constipation
Gerardo Calderon, Andres Acosta
SECTION E: Oncology
88 Cancer and Aging: General Principles
Carolyn J. Presley, Harvey Jay Cohen, Mina S. Sedrak
89 Breast Disease
Mina S. Sedrak, Hyman B. Muss
90 Prostate Cancer
Liang Dong, Mark C. Markowski, Kenneth J. Pienta
91 Lung Cancer
Asrar Alahmadi, Ajeet Gajra, Carolyn J. Presley
92 Gastrointestinal Malignancies
Ryan D. Nipp, Nadine J. McCleary
93 Skin Cancer
Shreya A. Sreekantaswamy, Suzanne Olbricht, Jonathan Weiss, Daniel C. Butler
SECTION F: Hematology
94 Aging of the Hematopoietic System and Anemia
Jiasheng Wang, Changsu Park, Jino Park, Robert Kalayjian, William Tse
95 Hematologic Malignancies (Leukemia/Lymphoma) and Plasma Cell Disorders
Anita J. Kumar, Tanya M. Wildes, Heidi D. Klepin, Bayard L. Powell
96 Coagulation Disorders
Ming Y. Lim
SECTION G: Endocrinology and Metabolism
97 Aging of the Endocrine System and Non-Thyroid Endocrine Disorders
Bradley D. Anawalt, Alvin M. Matsumoto
98 Thyroid Diseases
Anne R. Cappola
99 Diabetes Mellitus
Pearl G. Lee, Jeffrey B. Halter
SECTION H: Rheumatology
100 Myopathies, Polymyalgia Rheumatica, and Giant Cell Arteritis
Vivek Nagaraja
101 Rheumatoid Arthritis and Other Autoimmune Diseases
Jiha Lee, Raymond Yung
102 Back Pain and Spinal Stenosis
Owoicho Adogwa, Una E. Makris, M. Carrington Reid
103 Fibromyalgia and Myofascial Pain Syndromes
Cheryl D. Bernstein, Simge Yonter, Aishwarya Pradeep, Jay P. Shah, Debra K. Weiner
SECTION I: Infectious Diseases
104 Infection and Appropriate Antimicrobial Selection
Kevin P. High
105 Bacterial Pneumonia and Tuberculosis
Juan González del Castillo, Francisco Javier Martín Sánchez
106 Urinary Tract Infections
Muhammad S. Ashraf, Mandy L. Byers
107 Other Viruses: Human Immunodeficiency Virus Infection and Herpes Zoster
Kristine M. Erlandson, Kenneth Schmader
108 Influenza, COVID-19, and Other Respiratory Viruses
Lauren Hartman, H. Keipp Talbot
Index
Contributors
CHAPTER AUTHORS
Numbers in brackets refer to the chapters written or cowritten by the
contributor.
Andres Acosta, MD, PhD
Assistant Professor of Medicine
Division of Gastroenterology and Hepatology
Mayo Clinic
Rochester, Minnesota [87]
Owoicho Adogwa, MD, MPH
Associate Professor of Neurosurgery
Department of Neurological Surgery
University of Cincinnati School of Medicine
Cincinnati, Ohio [102]
Kathleen M. Akgün, MD, MS
Associate Professor of Medicine
Section of Pulmonary, Critical Care and Sleep Medicine
Yale University School of Medicine
New Haven, Connecticut
Director, Medical Intensive Care Unit
VA Connecticut Healthcare System
West Haven, Connecticut [81]
Asrar Alahmadi, MBBS MAS-CR
Division of Oncology, Department of Medicine
The Ohio State University Comprehensive Cancer Center
Columbus, Ohio [91]
Steven M. Albert, PhD, MS, FGSA, FAAN
Editor-in-Chief, Innovation in Aging
Professor and Hallen Chair
Department of Behavioral and Community Health Sciences
Graduate School of Public Health
University of Pittsburgh
Pittsburgh, Pennsylvania [4]
Cathy Alessi, MD
Director, Geriatric Research, Education and Clinical Center; VA Greater Los
Angeles Healthcare System
Professor, Department of Medicine
University of California, Los Angeles
Los Angeles, California [44]
Karen P. Alexander, MD
Professor of Medicine/Cardiology
Duke University and Duke Clinical Research Institute
Durham, North Carolina [74]
Bradley D. Anawalt, MD
Professor and Vice Chair
Department of Medicine
University of Washington School of Medicine
Chief of Medicine, University of Washington Medical Center
Seattle, Washington [97]
Hidenori Arai, MD, PhD
President
National Center for Geriatrics and Gerontology
Obu, Aichi, Japan [13]
Muhammad S. Ashraf, MBBS
Associate Professor
Division of Infectious Diseases, Department of Internal Medicine
University of Nebraska Medical Center
Omaha, Nebraska [106]
Tara Asokan, BS
Graduate Intern
The John A. Hartford Foundation
New York, New York [12]
Sanjay Asthana, MD, FACP
Associate Dean for Gerontology
Professor and Head, Division of Geriatrics and Gerontology
Director, NIA/NIH Wisconsin Alzheimer Disease
Research Center (ADRC)
Director, Madison VAMC Geriatric Research,
Education and Clinical Center (GRECC)
Duncan G. and Lottie H. Ballantine Endowed Chair in Geriatrics
Madison, Wisconsin [59]
Steven N. Austad, PhD
Protective Life Endowed Chair in Healthy Aging Research
Department of Biology
University of Alabama at Birmingham
Birmingham, Alabama [5]
Thilinie D. Bandaranayake, MBBS
Section of Infectious Diseases
Yale School of Medicine
New Haven, Connecticut [3]
Steven Barczi, MD FAASM
Director of Clinical Services, Division of Geriatrics, Department of
Medicine
Director of Clinical, Geri-PACT & GRECC Connect
Madison VA Geriatric Research, Education and Clinical Center
Professor of Medicine, Geriatrics/Sleep Medicine
University of Wisconsin-Madison School of Medicine and Public Health
Roxanne Bavarian, DMD
Orofacial Pain Fellow
Department of Oral and Maxillofacial Surgery
Massachusetts General Hospital
Boston, Massachusetts [68]
Christina L. Bell, MD, PhD
Clinical Associate Professor
Department of Geriatric Medicine
John A. Burns School of Medicine
University of Hawaii
Hawaii Permanente Medical Group
Honolulu, Hawaii [72]
Mark Benson, MD
Associate Professor
Division of Gastroenterology and Hepatology
University of Wisconsin
Amy Berman, BS, RN, LHD (hon), FAAN
Senior Program Officer
Cheryl D. Bernstein, MD
Associate Professor
Departments of Anesthesiology and Perioperative Medicine and Neurology
University of Pittsburgh School of Medicine
John Best, MD
Behavioral Neurology Clinical Fellow
Weill Institute for Neurosciences
Memory and Aging Center
University of California San Francisco
San Francisco, California [63]
Nirav R. Bhatt, MD
Assistant Professor of Neurology
Marcus Stroke and Neuroscience Center at Grady Memorial Hospital and
Emory University School of Medicine
Atlanta, Georgia [62]
Jawad Bilal, MD
Clinical Assistant Professor
Division of Rheumatology, Department of Internal Medicine
University of Arizona Arthritis
Tucson, Arizona [52]
Ellen F. Binder, MD
Professor of Medicine
Program Director, Geriatric Medicine Fellowship Program
Washington University in St. Louis, School of Medicine
St. Louis, Missouri [53]
Joyce M. Black, PhD, RN, FAAN
Florence Niedfelt Professor
College of Nursing
Caroline S. Blaum, MD, MS
Senior Research Scientist
National Committee for Quality Assurance
Washington, DC
Adjunct Professor Geriatric Medicine and Palliative Care
NYU Grossman School of Medicine
Laura Block, BS, BSN, RN
Nurse, PhD Student
School of Nursing
University of Wisconsin-Madison
Nicolaas I. Bohnen MD, PhD
Professor of Radiology (Division of Nuclear Medicine) and Neurology
University of Michigan
Staff Neurologist & GRECC Investigator
VA Ann Arbor Healthcare System
Ann Arbor, Michigan [61]
Alice F. Bonner, PhD, RN, FAAN
Adjunct Faculty and Director of Strategic Partnership
CAPABLE Program
Johns Hopkins School of Nursing
Baltimore, Maryland [16]
Cynthia Boyd, MD, MPH
Professor of Medicine, Epidemiology and Health Policy and Management
Division of Geriatric Medicine and Gerontology
Center for Transformative Geriatric Research
Johns Hopkins University School of Medicine
Jennifer S. Brach, PT, PhD, FAPTA
Professor
Department of Physical Therapy
Associate Dean for Faculty Affairs and Development, School of Health and
Rehabilitation Sciences
Francesca Brancati, BS
Amber K. Brooks, MD
Associate Professor, Department of Anesthesiology
Wake Forest University School of Medicine
Winston Salem, North Carolina [68]
Cynthia J. Brown, MD, MSPH, FACP
Charles V. Sanders, MD Endowed Chair in Internal Medicine
Professor and Chair
Louisiana State University Health Sciences Center
New Orleans, Louisiana [55]
William Bryson, MD, MPH
Assistant Clinical Professor
Department of Psychiatry and Behavioral Sciences
University of Washington
Nitin Budhwar, MD
Associate Professor of Internal Medicine
Division of General Internal Medicine
Section Chief Geriatric Medicine
University of New Mexico
Albuquerque, New Mexico [83]
Daniel C. Butler, MD
Assistant Professor, Director of UCSF Aging Skin and Geriatric
Dermatology Clinic
Department of Dermatology
University of California, San Francisco
Mandy L. Byers, MD, CMD
Assistant Professor
Division of Geriatrics, Gerontology, and Palliative Medicine, Department of
Internal Medicine
Joseph M. Calabrese, DMD
Clinical Professor
Director of Geriatric Dental Medicine
Associate Dean of Students
Henry M. Goldman School of Dental Medicine
Boston University
Gerardo Calderon, MD
Physician Resident
Indiana University School of Medicine
Indianapolis, Indiana [87]
Brook Calton, MD, MHS
Division of Palliative Medicine and Geriatrics
The Massachusetts General Hospital
Anne R. Cappola, MD, ScM
Division of Endocrinology, Diabetes, and Metabolism
Director, Penn Medical Communication Research Institute
Perelman School of Medicine at the University of Pennsylvania
Philadelphia, Pennsylvania [98]
Whitney L. Carlson, MD
Department of Psychiatry and Behavioral Sciences
University of Washington
Cynthia M. Carlsson, MD, MS
Division of Geriatrics and Gerontology
Christopher R. Carpenter, MD, MSc
Professor of Emergency Medicine
Department of Emergency Medicine
Washington University in St. Louis School of Medicine
Carol K. Chan, MBBCh, MSc
Department of Psychiatry
Case Western Reserve University School of Medicine
Cleveland, Ohio [60]
Elizabeth N. Chapman, MD
Associate Director, Education and Evaluation
Geriatrics Research Education and Clinical Center
William S. Middleton Memorial VA Medical Center
Madison, Wisconsin
Department of Medicine – Division of Geriatrics and Gerontology
Nathaniel A. Chin, MD
Brenna Cholerton, PhD
Senior Research Scientist/Neuropsychologist
Department of Pathology
Stanford University School of Medicine
Stanford, California [57]
Monica Christmas, MD, NCMP
Associate Professor
Department of Obstetrics and Gynecology
University of Chicago
Chicago, Illinois [35]
Jacqueline C. T. Close, MBBS, MD, FRCP, FRACP
Geriatrician and Conjoint Professor
Prince of Wales Clinical School
University of New South Wales
Sydney, Australia [13]
Harvey Jay Cohen, MD
Walter Kempner Professor of Medicine
Director Emeritus, Center for the Study of Aging and Human Development
Chair Emeritus, Department of Medicine
Duke University Medical Center
Jessica Colburn, MD
Division of Geriatric Medicine & Gerontology
Lisa Cooper, MD
Division of Aging
Brigham and Woman’s Hospital
Boston, Massachusetts
Department of Geriatric Medicine, Rabin Medical Center
Petach Tikva, Israel
Instructor of Medicine, Sackler Faculty of Medicine
Tel Aviv University
Tel Aviv, Israel [70]
Suzanne Craft, PhD
Director, Wake Forest Alzheimer’s Disease Research Center
Co-Director, Sticht Center on Healthy Aging and Alzheimer’s Prevention
Wake Forest School of Medicine
Winston-Salem, North Carolina [57]
Alfonso J. Cruz-Jentoft, MD, PhD
Jefe del Servicio de Geriatría
Hospital Universitario Ramón y Cajal (IRYCIS)
Madrid, Spain [49]
Juan González del Castillo, MD, PhD
Head of the Infectious Disease Group
The Spanish Emergency Medicine Society
Head of Emergency Department
Clínico San Carlos Hospital
Madrid, Spain [105]
Shannon Devlin, MD
Instructor of Medicine
Department of Internal Medicine, Division of Palliative Medicine
Washington University in St. Louis School of Medicine
Maryama Diaw, MPH
Graduate Intern
Nikola Dobrilovic, MD
Assistant Professor of Surgery
Boston University School of Medicine
Liang Dong, MD
Post-doctoral Research Fellow
Brady Urological Institute
Margaret A. Drickamer, MD
Professor, Division of Geriatric Medicine & Palliative Care Program
Medical Director, UNC Home Hospice & Hospice Home
School of Medicine
University of North Carolina at Chapel Hill
Chapel Hill, North Carolina [10]
Catherine E. DuBeau, MD
Section of General Internal Medicine – Geriatrics
Dartmouth-Hitchcock Medical Center
Lebanon, New Hampshire [38]
Ruth E. Dunkle, PhD
Wilbur J. Cohen Collegiate Professor of Social Work
School of Social Work
Gustavo Duque, MD, PhD, FRACP, FGSA
Geriatrician, Chair of Medicine and Director of the Australian Institute for
Musculoskeletal Science (AIMSS)
The University of Melbourne and Western Health
St Albans, Australia [51]
Katharina V. Echt, PhD
Associate Director Education and Evaluation
Veterans Affairs Birmingham/Atlanta Geriatric Research Education and
Clinical Center (GRECC)
Associate Professor
Department of Medicine, Geriatrics and Gerontology
Niloo M. Edwards, MD
Chief Cardiothoracic Surgery
Professor of Surgery
Boston University
Kristine M. Erlandson, MD
Associate Professor of Medicine and Epidemiology
Divisions of Infectious Diseases and Geriatric Medicine
University of Colorado—Anschutz Medical Campus
Aurora, Colorado [107]
Sara E. Espinoza, MD, MSc
Professor
Barshop Institute for Longevity and Aging Studies
University of Texas Health Science Center at San Antonio
Director, Geriatrics Research, Education and Clinical Center
South Texas Veterans Health Care System
San Antonio, Texas [40]
Timothy W. Farrell, MD, AGSF
Geriatrics Division Associate Chief for Age-Friendly Care
Spencer Fox Eccles School of Medicine at the University of Utah
Physician Investigator, VA Salt Lake City Geriatric Research, Education, and
Clinical Center
Salt Lake City, Utah [72]
Mizhgan Fatima, MBBS, FRACP
Geriatrician, Department of Geriatric Medicine, Western Health
Australian Institute for Musculoskeletal Science (AIMSS)
The University of Melbourne
St Albans, Australia [51]
Cynthia Felix, MD, PhD
Postdoctoral Associate
Department of Epidemiology
Luigi Ferrucci, MD, PhD
Scientific Director
National Institute on Aging
National Institutes of Health
Tamara G. Fong, MD, PhD
Associate Professor of Neurology, Harvard Medical School
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Laura Frain, MD, MPH
Director of Outpatient Geriatrics
Division of Aging
Brigham and Women’s Hospital
Instructor of Medicine, Harvard Medical School
Marilisa Franceschi, MD, PhD
Endoscopic Unit, Department of Medicine
Azienda ULSS 7 Pedemontana - Alto Vicentino Hospital
Via Garziere, Santorso (Vicenza), Italy [85]
Kaitlyn Fruin, MD
Resident Physician
Department of Internal Medicine
University of California Los Angeles
Los Angeles, California [35]
Terry Fulmer, PhD, RN, FAAN
President
Ajeet Gajra, MD, FACP
Clinical Professor
SUNY Upstate Medical University and Hematology-Oncology Associates of
Central New York
Syracuse, New York [91]
James E. Galvin, MD, MPH
Professor of Neurology
Chief, Division of Cognitive Neurology
Director, Comprehensive Center for Brain Health
University of Miami Miller School of Medicine
Miami, Florida [9]
Chandrika Garner, MD, FASE
Section on Cardiothoracic Anesthesiology
Department of Anesthesiology
Kathleen M. Gavin, PhD
Assistant Professor
Department of Medicine, Division of Geriatric Medicine
University of Colorado Anschutz Medical Campus
Sudeep S. Gill, MD, MSc, FRCPC
Associate Professor
Department of Medicine Division of Geriatric Medicine
Queen’s University
Kingston, Ontario, Canada [22]
Andrea Gilmore-Bykovskyi, PhD, RN
Deputy Director, Center for Health Disparities Research
Associate Professor, School of Nursing
University of Wisconsin
Madison, Wisconsin [6] [18]
Carey E. Gleason, PhD
Parag Goyal, MD, MSc
Director, Heart Failure with Preserved Ejection Fraction and Cardiac
Amyloidosis Program
Weill Cornell Medicine
New York Presbyterian Hospital
Len Gray, MBBS, MMed, PhD, FRACP, FACHSM, FAAG, FANZSGM
Director: Centre for Health Services Research
Faculty of Medicine
The University of Queensland
Brisbane, Australia [13]
Leanne Groban, MS, MD
Professor
Department of Anesthesiology
Matthew E. Growdon, MD, MPH
Geriatrician and Aging Research Fellow
Division of Geriatrics
Daniel Guidot, MD, MPH
Fellow
Division of Pulmonary and Critical Care
Jerry H. Gurwitz, MD
The Dr. John Meyers Professor of Primary Care Medicine
Professor of Medicine, Family Medicine and Community Health, and
Population & Quantitative Health Sciences
Chief, Division of Geriatric Medicine
Executive Director, Meyers Health Care Institute
UMass Chan Medical School
Worcester, Massachusetts [22]
Karen E. Hall, MD, PhD
Emeritus Professor of Medicine
Division of Geriatric and Palliative Medicine
Jeffrey B. Halter, MD
Professor Emeritus
Parkway Visiting Professor in Geriatrics
Yong Loo Lin School of Medicine
National University of Singapore
Benjamin H. Han, MD, MPH
Assistant Professor
Division of Geriatrics, Gerontology, and Palliative Care
University of California, San Diego
La Jolla, California [23]
Lauren Hartman, MD
Assistant Professor
Gerontology and Geriatric Medicine
Atrium Health Wake Forest Baptist
Jennifer Hayashi, MD
Kevin P. High, MD, MS
President, Atrium Health—Wake Forest Baptist
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Kerry L. Hildreth, MD
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Graduate Intern
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Section Chief, Cardiology
Ula Hwang, MD, MPH
Professor, Vice Chair for Research
Department of Emergency Medicine
Core Physician Investigator
Geriatric Research, Education and Clinical Center
James J. Peters VA Medical Center
Bronx, New York [15]
Sharon K. Inouye, MD, MPH
Director, Aging Brain Center
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Senior Associate Dean for Healthy Aging and Senior Care
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UC San Diego Center for Healthy Aging
Distinguished Professor of Psychiatry and Neurosciences
Director, Sam and Rose Stein Institute for Research on Aging
Department of Psychiatry, University of California, San Diego
San Diego, California [66]
Jason M. Johanning, MD, MS
Professor of Surgery
Vice Chair for Surgery: Quality and Compliance
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Larry E. Johnson, MD, PhD
Central Arkansas Veterans Healthcare System
Associate Professor of Geriatrics, and Family and Preventive Medicine
University of Arkansas for Medical Sciences
Little Rock, Arkansas [30]
Theodore M. Johnson, II, MD, MPH, AGS-Fellow
Paul Seavey Chair and Chief, General Internal Medicine
Chair, Family and Preventive Medicine
Investigator, Birmingham/Atlanta VA GRECC
Judith A. Jones, DDS, MPH, DScD
Professor
University of Detroit Mercy School of Dentistry
Detroit, Michigan [32]
Jamie N. Justice, PhD
Assistant Professor
Section on Gerontology & Geriatrics, Sticht Center on Healthy Aging and
Alzheimer’s Prevention
Robert Kalayjian, MD
Department of Medicine, MetroHealth System
Jennifer M. Kapo, MD
Associate Professor of Palliative Medicine
Chief, Palliative Medicine, Palliative Care Program
Section of Palliative Medicine
Marshall B. Kapp, JD, MPH
Professor Emeritus
Center for Innovative Collaboration in Medicine and Law
Florida State University, College of Medicine and College of Law
Tallahassee, Florida [26]
Jay Kayser, MSW, LCSW
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School of Social Work and Department of Developmental Psychology
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Harvard Medical School
Hinda and Arthur Marcus Institute for Aging Research
Hebrew SeniorLife
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Associate Dean for Social Health Sciences and Programs
Director, UW Center for Health Disparities Research
Madison, Wisconsin [6] [18]
Daniel Kirsch, BA
MD/PhD Candidate
Jamaica Plain VA Medical Center
Dalane W. Kitzman, MD
Professor of Internal Medicine
Sections on Cardiovascular Medicine and Geriatrics/Gerontology
Heidi D. Klepin, MD, MS
Professor
Section on Hematology and Oncology
Nway Le Ko Ko, MD
Cardiology Fellow
Mayo Clinic College of Medicine
Department of Cardiovascular Diseases
Mayo Clinic Arizona
Phoenix, Arizona [77]
Vikas Kotagal, MD, MS
Associate Professor of Neurology
Staff Neurologist & GRECC Investigator
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UCSF and San Francisco VA Medical Center
George A. Kuchel, MD CM, AGSF
Professor and Travelers Chair in Geriatrics and Gerontology
Director, UConn Center on Aging
Chief, Geriatric Medicine
University of Connecticut and UConn Health
Farmington, Connecticut [39] [40]
Alexis Kuerbis, LCSW, PhD
Associate Professor
Silberman School of Social Work
Hunter College
Jeffrey T. Kullgren, MD, MS, MPH
Research Scientist
VA Center for Clinical Management Research
Associate Professor
Division of General Medicine
Anita J. Kumar, MD, MSc
Assistant Professor
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Hiroko Kunitake, MD, MPH
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Department of Surgery
Massachusetts General Hospital
C. Kent Kwoh, MD
Professor of Medicine and Medical Imaging
Division of Rheumatology, Department of Medicine, and University of
Arizona Arthritis Center
University of Arizona College of Medicine and Banner University Medical
Center
Mark S. Lachs, MD, MPH
Psaty Distinguished Professor of Medicine
Weill Cornell Medicine
Director of Geriatrics
New York Presbyterian Health System
Jeffrey Lam, BA
Warren Alpert Medical School of Brown University
Providence, Rhode Island [66]
Ellen E. Lee, MD
Assistant Professor of Psychiatry
Staff Psychiatrist
Veterans Affairs San Diego Healthcare System
San Diego, California [66]
Jiha Lee, MD, MHS
Assistant Professor
Patty J. Lee, MD
Professor of Medicine, Cell Biology & Pathology
Division of Pulmonary, Allergy and Critical Care
Duke University School of Medicine
Durham, North Carolina [80] [81]
Pearl G. Lee, MD
Geriatric Research Education and Clinical Center (GRECC)
Sei J. Lee, MD, MAS
UCSF and San Francisco VA Medical Center
Bruce Leff, MD
Bernardo Liberato, MD
Assistant Professor of Neurology
Marcus Stroke and Neuroscience Center at Grady Memorial Hospital and
Ming Y. Lim, M.B.B.Chir
Associate Professor
Division of Hematology and Hematologic Malignancies
University of Utah
Salt Lake City, Utah [96]
Stacy Tessler Lindau, MD, MAPP
Professor
Departments of Obstetrics and Gynecology and Medicine-Geriatrics and
Palliative Medicine
University of Chicago
Chicago, Illinois [35]
Lewis A. Lipsitz, MD
Professor of Medicine, Harvard Medical School
Chief Academic Officer and Director, Marcus Institute for Aging Research,
Hebrew Senior Life
Chief, Division of Gerontology, Beth Israel Deaconess Medical Center
Editor-in-Chief, Journal of Gerontology Medical Sciences
Julia Loewenthal, MD
Instructor in Medicine
Division of Aging
Brigham and Women’s Hospital
David B. Lombard, MD, PhD
Associate Professor
Stephen R. Lord, PhD, DSc
Professor
Senior Principal Research Fellow
Falls Balance and Injury Research Centre
Neuroscience Research Australia
Michael R. Lucey, MD, FRCPI, FAASLD
Chief, Division of Gastroenterology and Hepatology
Co-Medical Director, UW Digestive Health Center
Medical Director, UW Liver Transplant Program
Hillary D. Lum, MD, PhD
Division of Geriatric Medicine
University of Colorado School of Medicine
Constantine G. Lyketsos, MD, MHS
Chair, Department of Psychiatry and Behavioral Sciences, Johns Hopkins
Bayview
Elizabeth Plank Althouse Professor, Johns Hopkins University
Director, Richman Family Precision Medicine Center of Excellence in
Alzheimer’s Disease
Director, Johns Hopkins Memory and Alzheimer’s Treatment Center
Tanya Mailhot, RN, PhD
Researcher, Montreal Heart Institute Research Center
Assistant Professor, Faculty of Nursing
Université de Montréal
Montréal, Canada [58]
Una E. Makris, MD, MSc
Assistant Professor
Department of Internal Medicine, Division of Rheumatic Diseases
UT Southwestern Medical Center
Dallas, Texas [102]
Preeti N. Malani, MD, MSJ
Chief Health Officer
Michael L. Malone, MD
Medical Director, Aurora Senior Services & Aurora at Home
Geriatrics Fellowship Director, Aurora Sinai Medical Center
Clinical Adjunct Professor of Medicine
University of Wisconsin School of Medicine & Public Health
Co-Editor, Journal of Geriatric Emergency Medicine (JGEM)
Section Editor, Journal of the American Geriatrics Society (JAGS)
Models of Geriatric Care, Quality Improvement and Program Dissemination
Milwaukee, Wisconsin [14]
Mark C. Markowski, MD, PhD
Assistant Professor
Department of Oncology
Michelle M. Marrero, MD
Instructor of Neurology
Division of Cognitive Neurology
Department of Neurology
University of Miami Miller School of Medicine
Miami, Florida [9]
Finbarr C. Martin, MD, MSc, FRCP
Emeritus Geriatrician and Professor of Medical Gerontology
Population Health Sciences, Faculty of Life Sciences and Medicine
King’s College London
London, United Kingdom [13]
Kedar S. Mate, MD
President and Chief Executive Officer
Institute for Healthcare Improvement
Boston, Massachusetts
Weill Cornell Medical College
Daniel D. Matlock, MD, MPH
Division Head (Interim), Division of Geriatrics
Director, Colorado Program for Patient Centered Decisions
University of Colorado School of Medicine
Alvin M. Matsumoto, MD
Professor Emeritus, Department of Medicine
Division of Gerontology and Geriatric Medicine
University of Washington School of Medicine
Clinical Investigator, Geriatric Research, Education and Clinical Center
VA Puget Sound Health Care System
Seattle, Washington [37] [97]
Mathew S. Maurer, MD
Arnold and Arlene Goldstein Professor of Cardiology
Director, Clinical Cardiovascular Research Laboratory for the Elderly
Columbia University Irving Medical Center
New York Presbyterian Hospital
Nadine J. McCleary, MD, MPH
Assistant Professor of Medicine, Harvard Medical School
Senior Physician, Dana-Farber Cancer Institute
Shelley R. McDonald, DO, PhD
Associate Professor
Ann C. McKee, MD
William Fairfield Warren Distinguished Professor of Neurology and
Pathology
Director of Neuropathology VA Boston
Director of the BU Chronic Traumatic Encephalopathy (CTE) Center
Simon Mears, MD, PhD
Professor of Orthopedic Surgery
University of Arkansas for Medical Sciences
Little Rock, Arkansas [53]
Darshan H. Mehta, MD, MPH
Education Director
Osher Center for Integrative Medicine
Brigham and Women’s Hospital and Harvard Medical School
Jasmine C. Menant, PhD
Research Fellow
Falls Balance and Injury Research Centre
Neuroscience Research Australia
David J. Meyers, PhD, MPH
Assistant Professor
Department of Health Services, Policy, and Practice
Brown University School of Public Health
Providence, Rhode Island [19]
Christine Miaskowski, RN, PhD, FAAN
Professor of Physiological Nursing
School of Nursing, University of California
University of California San Francisco School of Nursing
Bruce Miller, MD
A.W. and Mary Margaret Clausen Distinguished Professor of Neurology
Director, Memory and Aging Center
Richard A. Miller, MD, PhD
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Alison A. Moore, MD, MPH, FACP, AGSF
Larry L. Hillblom Chair in Geriatric Medicine
Chief, Division of Geriatrics, Gerontology, and Palliative Care
La Jolla, California [23]
Kerrie L. Moreau, PhD
Professor
Department of Medicine, Division of Geriatrics
Hyman B. Muss, MD
Mary Jones Hudson Distinguished Professor of Geriatric Oncology
Director of Geriatric Oncology, Lineberger Comprehensive Cancer Center
University of North Carolina
Chapel Hill, North Carolina [89]
Vivek Nagaraja, MD
Division of Rheumatology
Aanand D. Naik, MD
Professor and Nancy P. & Vincent F. Guinee, MD Distinguished Chair
Department of Management, Policy and Community Health, University of
Texas School of Public Health
Executive Director, UTHealth Institute on Aging
University of Texas Health Science Center
Houston, Texas
Investigator, Houston Center for Innovations in Quality, Effectiveness and
Safety
Michael E. DeBakey VA Medical Center
Houston, Texas [25]
Abhijit S. Naik, MD, MPH, FASN
Assistant Professor of Internal Medicine
Division of Nephrology
Michael G. Nanna, MD, MHS
Assistant Professor of Medicine/Cardiology
Anne B. Newman, MD, MPH
Distinguished Professor and Chair
Department of Epidemiology, Graduate School of Public Health
John C. Newman, MD, PhD
Assistant Professor
Buck Institute for Research on Aging
University of California San Francisco
Novato, California [40]
Ryan D. Nipp, MD
Medical Oncology at Stephenson Cancer Center
University of Oklahoma Health Sciences Center
Oklahoma City, Oklahoma [92]
Stephanie Nothelle, MD
Assistant Professor of Medicine and Health Policy and Management
Division of Geriatric Medicine and Gerontology
Center for Transformative Geriatric Research
Michele R. Obert, MD
Internal Medicine Resident
University of Arizona
Michelle C. Odden, PhD
Associate Professor
Department of Epidemiology and Population Health
Stanford School of Medicine
Stanford University
Stanford, California [2]
Suzanne Olbricht, MD
Chief of Dermatology
Beth Israel Dermatology, Beth Israel Deaconess Medical Center
Associate Professor of Dermatology
Christopher D. Ortengren, MD
Senior Resident Physician
Department of Surgery
Section of Urology
Dartmouth-Hitchcock Medical Center
One Medical Center Drive
Lebanon, New Hampshire [38]
Joseph G. Ouslander, MD
Professor of Geriatric Medicine
Senior Advisor to the Dean for Geriatrics
Charles E. Schmidt College of Medicine
Professor (Courtesy), Christine E. Lynn College of Nursing
Florida Atlantic University
Boca Raton, Florida [16] [19]
Ambarish Pandey, MD, MSCS
Assistant Professor of Internal Medicine
Division of Cardiology
University of Texas Southwestern Medical Center
Dallas, Texas [73]
Changsu Park, MD
Jino Park, PhD
Assistant Professor
Division of Hematology/Oncology
Sanjeevkumar R. Patel, MD, MS
Associate Professor
Leslie Pelton, MPA
Vice President
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University of California, Berkeley
School of Social Welfare
Berkeley, California [20]
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Brady Urological Institute
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Chair, Division of Geriatric Medicine & Gerontology
Robert and Arlene Kogod Professor of Geriatric Medicine
Mayo Clinic College of Medicine
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Director Geriatrics Unit
Director Department Geriatric Care, OrthoGeriatrics and Rehabilitation
Galliera Hospital
Genova, Italy
Full Professor in Geriatrics
Department of Interdisciplinary Medicine
University of Bari “Aldo Moro”
Bari, Italy [85]
Scott D. Pletcher, PhD
Professor
Department of Molecular and Integrative Physiology
Jennifer D. Possick, MD
Section of Pulmonary, Critical Care and Sleep Medicine
Medical Director, Winchester Center for Lung Disease and Director, Post-
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Yale-New Haven Hospital
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University of Alabama at Birmingham
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Aishwarya Pradeep, BS
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majority are fluviatile, one group (Neritodryas) actually occurring in
the Philippines on trees of some height, at a distance of a quarter of
a mile from any water. Navicella is a still further modified form of
Neritina, occurring only on wet rocks, branches, etc., in non-tidal
streams (Fig. 13).
Fig. 13.—Illustrating the development of the fresh-water genus

Navicella, through the brackish-water Neritina, from the marine
Nerita, with corresponding changes in the operculum. 1.
Nerita; 2, 3. Neritina; 4. Neritina, intermediate form; 5, 6.
Navicella.
The great family of the Melaniidae, which occurs in the rivers of
warm countries all over the world, and that of the Pleuroceridae,
which is confined to North America, are, in all probability, derived
from some form or forms of Cerithium. The origin of the Paludinidae,
Valvatidae, and Ampullariidae is more doubtful. Their migration from
the sea was probably of an early date, since the first traces of all
three appear in the lower Cretaceous, while Melaniidae are not
known until Tertiary times. Ampullaria, however, shows distinct signs
of relationship to Natica, while the affinities of Paludina and Valvata
cannot as yet be approximately affirmed.
(2) Pulmonata.—Intermediate between the essentially fresh-water
and the essentially marine species come the group sometimes
known as Gehydrophila, consisting of the two families Auriculidae
and Otinidae. These may be regarded as Mollusca which, though
definitely removed from all marine species by the development of a
true lung or lung cavity in the place of a gill, have yet never become,
in respect of habitat, genuine fresh-water species. Like Potamides,
they haunt salt marshes, mangrove swamps, and the region about
high-water mark. In some cases (Otina, Melampus, Pedipes) they
live on rocks which are moistened, or even bathed by the spray, in
others (Cassidula, Auricula) they are immersed in some depth of
brackish water at high tide, in others again (Scarabus) they are more
definitely terrestrial, and live under dead leaves in woods at some
little distance from water. Indeed one genus of diminutive size
(Carychium) has completely abandoned the neighbourhood of the
sea, and inhabits swampy ground almost all over the world.
Fig. 14.—Examples of the Auriculidae: A,
Auricula Judae Lam., Borneo; B, Scarabus
Lessoni Blainv., E. Indies; C, Cassidula
mustelina Desh., N. Zealand; D, Melampus
castaneus Mühlf., S. Pacific; E, Pedipes
quadridens Pfr., Jamaica.
Fig. 15.—An example of
Amphibola (avellana
Chem.), the only true
Pulmonate which
possesses an
operculum.
To this same section Gehydrophila have been assigned two
remarkable forms of air-breathing “limpet,” Siphonaria and Gadinia
(see page 151), and the aberrant Amphibola, a unique instance of a
true operculated pulmonate. Siphonaria possesses a pulmonary
cavity as well as a gill, while Gadinia and Amphibola are exclusively
air-breathing. Siphonaria lives on rocks at or above high-water mark,
Gadinia between tide marks, Amphibola (Fig. 15) in brackish water
at the estuaries of rivers, half buried in the sand. There can be little
doubt that all these are marine forms which are gradually becoming
accustomed to a terrestrial existence. In Gadinia and Amphibola the
process is so far complete that they have exchanged gills for a
pulmonary cavity, while in Siphonaria we have an intermediate stage
in which both organs exist together. A curious parallel to this is found
in the case of Ampullaria, which is furnished with two gills and a
pulmonary chamber, and breathes indifferently air and water. It is a
little remarkable that Siphonaria, which lives at a higher tide level
than Gadinia, should retain the gill, while Gadinia has lost it.
The ultimate affinities of the essentially fresh-water groups,
Limnaea, Physa, Chilina, cannot be precisely affirmed. The form of
shell in Latia, Gundlachia, and perhaps Ancylus, may suggest to
some a connexion with the Otinidae, and in Chilina, a similar
connexion with the Auriculidae. But, in a question of derivation,
similarities of shell alone are of little value. It is not a little
remarkable, for instance, that we should find a simple patelliform
shell in genera so completely distinct from one another in all
anatomical essentials as Ancylus, Patella, Siphonaria, Propilidium,
Hipponyx, Cocculina, and Umbrella.
Some recent authors, on grounds of general organisation, regard
the Limnaeidae and their allies as Opisthobranchs adapted to an
aerial life. It is held[24] that the Nudibranchiate Opisthobranchs have
given birth to the Pulmonata Stylommatophora or land snails, and
the Tectibranchiate Opisthobranchs to the Pulmonata
Basommatophora or fresh-water snails. Such a view seems at first
sight open to some objection from other views than those which deal
simply with anatomy. The Opisthobranchiata are not, to any marked
extent, littoral genera, nor do they specially haunt the mouths of
rivers. On the contrary, they inhabit, as a rule, only the very lowest
part of the littoral zone, and are seldom found, except where the
water is purely salt. In other cases, when the derivation of land or
fresh-water genera is fairly well established, intermediate forms
persist, which indicate, with more or less clearness, the lines along
which modification has proceeded. It has, however, recently been
shown that Siphonaria[25] and Gadinia,[26] which have, as has been
already mentioned, hitherto been classified as Pulmonata, are in
reality modified forms of Opisthobranchiata, which are in process of
adaptation to a life partly marine, partly on land. They may therefore
be regarded as supplying the link, hitherto missing, between the land
Pulmonata and the marine groups from one or other of which the
latter must have been derived. The general consensus of recent
opinion inclines towards accepting these views, some writers[27]
being content to regard the Pulmonata, as a whole, as derived from
the Tectibranchiate Opisthobranchs, while others[28] go further and
regard the Stylommatophora as derived directly from the
Basommatophora.
Origin of the Land Fauna

Gasteropoda.—(1) Operculate. On a priori grounds, one might
predict a double origin for land operculates. Marine species might be
imagined to accustom themselves to a terrestrial existence, after a
period, more or less prolonged, of littoral probation. Or again, fresh-
water species, themselves ultimately derived from the sea, might
submit to a similar transformation, after a preliminary or intermediate
stage of life on mudbanks, wet swamps, branches overhanging the
water, etc. Two great families in this group, and two only, seem to
have undergone these transformations, the Littorinidae and the
Neritidae. The derivation of almost all existing land operculates may
be referred to one or other of these groups.
Fig. 16.—Two rows of the radula of Littorina littorea L., × 72.

The power of the Littorinidae to live for days or even weeks
without being moistened by the sea may be verified by the most
casual observer. In the tropics this power seems even greater than
on our own shores. I have seen, in various parts of Jamaica, Littorina
muricata living at the top of low cliffs among grass and herbage. At
Panama I have taken three large species of Littorina (varia, fasciata,
pulchra), on trees at and above high-water mark. Cases have been
recorded in which a number of L. muricata, collected and put aside,
have lived for three months, and L. irrorata for four months.[29]
These facts are significant, when we know that the land operculates
almost certainly originated in a tropical climate.
The Cyclophoridae, Cyclostomatidae, and Aciculidae, which, as
contrasted with the other land operculates, form one group, have
very close relations, particularly in the length and formation of the
radula, or lingual ribbon, with the Littorinidae.
Fig. 17.—Two rows of the radula of Cyclophorus sp., India, ×

40.
On the other hand, the Helicinidae, Hydrocenidae, and
Proserpinidae are equally closely related to Neritina. The
Proserpinidae (restricted to the Greater Antilles, Central America and
Venezuela) may perhaps be regarded as the ultimate term of the
series. They have lost the characteristic operculum, which in their
case is replaced by a number of folds or lamellae in the interior of
the shell. It has already been noticed how one group of Neritina
(Neritodryas) occurs normally out of the water. This group furnishes
a link between the fresh-water and land forms. It is interesting to
notice that here we have the most perfect sequence of derivatives;
Nerita in the main a purely marine form, with certain species
occurring also in brackish water; Neritina in the main fresh-water, but
some species occurring on the muddy shore, others on dry land;
Helicina the developed land form; and finally Proserpina, an aberrant
derivative which has lost the operculum.[30]
Fig. 18.—A, Neritina reticularis Sowb., Calcutta (brackish
water); B, Helicina neritella Lam., Jamaica (land); C,
Proserpina (Ceres) eolina Ducl., Central America (land).
Gasteropoda.—(2) Pulmonata. The origin of these, the bulk of
the land fauna, must at present be regarded as a problem not yet
finally solved. Some authorities, as we have seen, regard them as
derived from the Nudibranchiate, others, probably more correctly,
from the Tectibranchiate Opisthobranchs.
The first known members of the land Pulmonata (Pupa [?],
Hyalinia) are from the Carboniferous of North America. Similar but
new forms appear in the Cretaceous, from which time to the present
we have an unbroken series. The characteristically modern forms,
according to Simroth,[31] are Helices with thick shells. According to
the same author, Vitrina and Hyalinia are ancestral types, which give
origin not only to many modern genera with shells, but to many shell-
less genera also, e.g. Testacella is probably derived through
Daudebardia from Hyalinia, while from Vitrina came Limax and
Amalia. A consideration of the radulae of the genera concerned
certainly tends in favour of these views.
Godwin-Austen, speaking generally, considers[32] genera of land
Pulmonata with strongly developed mantle-lobes and rudimentary
shell as more advanced in development than genera in which the
shell is large and covers all or nearly all the animal.
CHAPTER II
LAND AND FRESH-WATER MOLLUSCA, THEIR HABITS AND GENERAL
ECONOMY
The majority of the Land Mollusca are probably more sensitive

than is usually believed. The humidity of the air must affect the
surface of their skin to a considerable extent. Every one has noticed
how the snails ‘come out’ on a damp evening, especially after rain.
As a rule, they wait till rain is over, probably objecting to the patter of
the drops upon their delicate tentacles. Snails kept in captivity under
a bell-glass are acutely sensitive of a damp atmosphere, and will
bestir themselves after rain just as if they were in the open air.
Certain Helices which are accustomed to live in moist places, will
find their way to water, if removed from their usual haunts. A case is
recorded[33] of a specimen of H. arbustorum, kept in a kitchen, which
used to find its way directly under the cold water tap, and appeared
to enjoy the luxury of a douche. How delicately the conditions of life
are balanced in some of these creatures is seen in the case of
Omalonyx, a genus akin to Succinea, which is found in Brazil and
the northern parts of South America. It lives creeping on plants which
overhang the margin of water, but perishes equally, if placed in the
water itself, or removed to a distance from it for any length of time.
[34]
Endurance of Heat and Cold.—The Mollusca are capable, at

least as far as some species are concerned, of enduring severe
extremes both of cold and heat. The most northern pulmonate yet
observed is a fresh-water species, Physa (Aplecta) hypnorum L. This
hardy mollusc, whose shell is so fragile as to need most careful
handling, has been noticed on the peninsula of Taimyr, North Siberia,
in 73° 30’ N. lat, a region whose mean annual temperature is below
10° F. with a range of from 40° F. in July to -30° F. in January.
It is well known that the Limnaeidae, and probably most fresh-
water Mollusca of sub-temperate regions, can continue to live not
merely under, but enveloped in ice, and themselves frozen hard.
Garnier relates[35] that, during the winter of 1829–30, some large
Limnaea auricularia, which had been placed in a small basin, were
frozen into a solid mass, experiencing a cold of -2° F. He supposed
they were dead, but, to his surprise, when the basin thawed, the
Limnaea gradually revived. Paludina vivipara and Anodonta anatina
have been known to resist a temperature of 23° F., and the former
has produced young shortly after being thawed out of the ice.[36] As
far north as Bodø in Norway (67° 37’ N. lat., well within the Arctic
circle) there are found no less than fourteen species of terrestrial
Mollusca, among them being Balea perversa and Clausilia rugosa.
[37]
Vitrina is one of our most hardy molluscs, and may be observed

crawling on bright mornings over the frost-covered leaves of a wood
or copse. V. glacialis is said by Charpentier to live in the Alps at a
height where the stones are covered with snow from nine to ten
months of the year. Many of the Hyaliniae are very hardy. Arion, in
spite of having no external shell to protect it, is apparently less
affected by the cold than Helix, and does not commence hibernation
till a later period in the autumn. The operculate land Mollusca, in
spite of the protection which their operculum may be supposed to
afford, are exceedingly sensitive to cold, and the whole group is
without doubt a product of tropical or semi-tropical regions (see map
at frontispiece). A species of Helicina which inhabits the southern
States of North America has been known to be almost exterminated
from certain districts by the occurrence of an unusually severe
winter.
One of the highest altitudes at which a land shell is known to live
appears to be the Liti Pass (Himalayas, 14,000 ft.). At this enormous
altitude, two species of Buliminus (arcuatus Hutt. and nivicola Bens.)
live on juniper bushes among patches of snow. An Anadenus is said
to have been found in a similar locality at 15,000 ft., while Limnaea
Hookeri has been taken from over 16,400 ft. in Landour. In the
Andes of Peru and Bolivia, five species of Bulimulus, one of Pupa,
and one of Limax occur at an elevation of 10,500 to 15,000 ft.
Several fresh-water Mollusca inhabit Lake Titicaca, which stands at a
height of 12,550 ft. in the Bolivian table-land.
In certain parts of the desert of Algeria, where there is not a trace
of vegetation to be seen, and the temperature at mid-day is 110° F.,
the ground is sometimes so covered with Helix lactea as to appear
perfectly white. Dr. F. H. H. Guillemard has told me that he noticed,
in somewhat similar surroundings between Fez and Tangier, H.
pisana in such extraordinary abundance that they hung from the low
scrub in bunches the size of a man’s two fists. It is singular that
Mollusca should live, and not only live, but flourish, in localities
apparently so unpromising. Shells which occur in the Algerian
Sahara are actually larger and altogether finer than the ordinary
European form of the same species. In order to protect themselves
to some extent against the scorching heat and consequent
evaporation, desert species are frequently modified in one of two
ways; the shell becomes either white or a light dusky brown, as in
the familiar Helix desertorum, or else it gains immensely in
thickness. Specimens of H. pomatia, recently procured from Fez, are
of extraordinary thickness as compared with forms from our own
chalk downs of Kent and Surrey.
Fresh-water Mollusca are frequently found inhabiting hot springs.
Thus Neritina fluviatilis lives at Bagnères de Bigorre in water at about
68° F. In another hot spring in the eastern Pyrenees a Bithynia lives
at a temperature of over 73° F.; while Blainville mentions another
case of a Bithynia living in water at 122° F.
Hibernation and Aestivation.—As autumn begins to draw on,
and the first frosts to nip vegetation, terrestrial species retire beneath
stones, into cracks in old walls, holes in tree trunks, deep fissures in
rocks, and nooks and crannies of every kind, or else bury
themselves deeply in the earth or in moss and heaps of leaves. They
thus commence their period of hibernation, which varies in length
according to the duration of winter. Frequently masses of Helices
may be found attached to one another, probably not so much for the
sake of warmth, for their temperature is but low, as to share the
comforts of a cosy retreat in common. Slugs generally hibernate
alone, excavating a sort of nest in the earth, in which they encyst
themselves, contracting their bodies until they are almost round, and
secreting a covering of their own slime. The Helices usually close up
the mouth of their shell by the formation of a membranous or chalky
epiphragm, which will be further described below. Both snails and
slugs take care to be in good condition at the time their winter sleep
begins, and for this reason the former are said to be most esteemed
by foreign epicures if captured just at this period.[38]
During hibernation, the action of the heart in land Pulmonata
ceases almost entirely. This appears to be directly due to the effect
of cold. Mr. C. Ashford has related[39] some interesting experiments
made upon H. hortensis and Hyal. cellaria, with the view of
ascertaining the effect of cold upon their pulsations. His observations
may be tabulated as follows:—
Number of pulsations per minute
Helix hortensis Hyal. cellaria At degrees Fahr.
22 21 52°
14 12 44°
10 11 38°
4 9 30°
At low temperatures the character, as well as the number of the
pulsations changed; they became imperfect and intermittent,
although exceptionally at 31° F. a H. rufescens gave five or six
pulsations a minute, very full and deliberate. The result of taking the
Hyalinia suddenly into the heat of a greenhouse was to bring on
palpitations. Further experiments resulted in evidence of a similar
kind. Hyal. radiatula, placed upon a deal table in a room, showed 52
pulsations per minute at 62° F. Placed upon the palm of the hand,
the action soon rose to 108. Hyal. alliaria, similarly treated, rose from
72 pulsations to 110. Floated upon water, the action of the heart of
the latter suddenly fell to 29.
Fresh-water Pulmonata do not appear to hibernate. Unio and
Anodonta, however, bury themselves more deeply in the mud, and
Dreissensia casts off its byssus and retires under the mud in deeper
water.[40] Limnaea and Planorbis have often been noticed to crawl
about under the lower surface of a thick coating of ice. In periods of
prolonged drought, when the water in the ponds dries up, the
majority of genera bury themselves in the mud. I have known
Limnaea peregra bury itself three inches deep, when surprised by a
sudden fall of the water in the ditch on Coe Fen, behind Peterhouse,
Cambridge. Physa hypnorum frequents by preference ditches which
dry up in summer, as does also Planorbis spirorbis, the latter often
forming a sort of epiphragm against evaporation. Ancylus has been
observed to spend the whole winter out of water, and P. spirorbis has
been noticed alive after four months’ desiccation.[41]
True aestivation, however, occurs mainly in the tropics, where
there is no winter, but only a period when it is not quite so hot as the
rest of the year, or on a coast like the Mediterranean, which is
subject to sudden and severe heat. This period is usually rainless,
and the heat is therefore a dry heat. At this season, which may last
for three or four months, most of the land Mollusca enter upon a
period of inaction, either burying themselves deeply in the ground, or
else permanently attaching themselves to the stalks of grass and
other herbage, or the under sides of rocks. For instance, the large
and beautifully painted Orthalicus, Corona, and Porphyrobaphe,
which inhabit Brazil, Ecuador, and eastern Peru, bury themselves
deeply in the ground during the dry season, while in the rains they
climb to the topmost branches of the great forest trees.[42] Thus it
may well happen that a visitor to a tropical island, Ceylon for
instance, or one of the Greater Antilles, if he times his visit to
coincide with the rainless season, may be grievously disappointed at
what seems its unaccountable poverty in land Mollusca. But as soon
as the weather breaks, and the moisture penetrates their retreats,
every bush and every stone, in favoured localities, will be alive with
interesting species.
The Epiphragm.—A considerable number of the land Pulmonata
(and a very few of the fresh-water) possess the power of closing the
aperture of their shell by means of what is known as an epiphragm
or covering of hardened mucus. This epiphragm is habitually formed
by certain species during hibernation or aestivation, or even during
shorter periods of inactivity and retirement, the object being, either to
check evaporation of the moisture of the body, or to secure the
animal against the cold by retaining a thin layer of slightly warm air
immediately within the aperture of the shell.
The epiphragm differs widely in character in different species,
sometimes (Clausilia, Pupa, Planorbis) consisting of the merest
pellicle of transparent membrane, while at others (Helix aperta, H.
pomatia) it is a thick chalky substance, with a considerable
admixture of carbonate of lime, with the consistency of a hardened
layer of plaster of Paris. Within these extremes every variety of
thickness, solidity, and transparency occurs. During long hibernation
several epiphragms are not unfrequently formed by the same
individual snail, one within the other, at gradually lessening
distances. The epiphragm thus performs, to a certain extent, the part
of an operculum, but it must be remembered that it differs radically
from an operculum physiologically, in being only a temporary
secretion, while the operculum is actually a living part of the animal.
The actual mode of formation of the epiphragm would seem to
differ in different species. According to Fischer,[43] the mollusc
withdraws into its shell, completely blocking all passage of air into
the interior, and closing the pulmonary orifice. Then, from the middle
part of the foot, which is held exactly at the same plane as the
aperture, is slowly secreted a transparent pellicle, which gradually
thickens, and in certain species becomes calcareous. Dr. Binney,
who kept a large number of Helix hortensis in confinement, had
frequently an opportunity of noticing the manner in which the
epiphragm was formed.[44] The aperture of the shell being upward,
and the collar of the animal having been brought to a level with it, a
quantity of gelatinous matter is thrown out [? where from]. The
pulmonary orifice is then opened, and a portion of the air within
suddenly ejected, with such force as to separate the viscid matter
from the collar, and to project it, like a bubble of air, from the
aperture. The animal then quickly withdraws farther into the shell,
and the pressure of the external air forces back the vesicle to a level
with the aperture, when it hardens and forms the epiphragm. In
some of the European species in which the gelatinous secretion
contains more carbonate of lime, solidification seems to take place at
the moment when the air is expelled, and the epiphragm in these is
in consequence strongly convex.
Thread-spinning.—A considerable number of fresh-water
Mollusca possess the power of stretching a thread, which is no more
than an exceedingly elongated piece of mucus, to the surface of the
water, and of using it as a means of locomotion. This thread bears
no analogy whatever to the fibrous byssus of certain bivalves, being
formed in an entirely different manner, without the need of a special
gland.
The threads are ‘spun’ by several species of Limnaea, Physa, and
Planorbis, by Bithynia tentaculata, and several of the Cycladidae.
They are anchored to the surface by a minute concavity at the upper
end, which appears to act like a small boat in keeping the thread
steady. The longest threads are those of the Physae, which have
been noticed to attain a length, in confinement, of 14 inches. They
are always spun in the ascent, and as a rule, when the animal
descends, it rolls the thread up and carries it down as it goes. A
single thread is never spun on the descent, but occasionally, when a
thread has become more or less of a permanence, it becomes
stronger by the addition of more mucus each time it is used, whether
for ascending or descending purposes. Cyclas cornea appears to be
an exception to the rule that threads are only spun on the ascent.
This species, which is particularly fond of crawling along the under
surface of the water, has been noticed to spin a thread half an inch in
length while on the surface, and to hang suspended from it for a
considerable time.
What the exact use of the thread may be, must to a certain extent
be matter of conjecture. The Limnaeidae are, in the great majority of
cases, compelled to make periodic visits to the surface in order to
inspire oxygen. It is also a favourite habit with them to float just
under the surface, or crawl about on its under side, perhaps in
pursuit of tiny vegetable organisms. Whatever may be the object of
an excursion to the surface, a taut thread will obviously be a nearer
way up than any other which is likely to present itself; indeed, without
this thread-spinning power, which insures a tolerably rapid arrival at
the surface, the animal might find itself asphyxiated, or at least
seriously inconvenienced, before it could succeed in taking in the
desired supply of oxygen. With the Cycladidae, which do not breathe
air, such an explanation is out of place; in their case the thread
seems to be a convenient means of resting in one position in the
intervals of the periods of active exercise to which several of the
species are so much addicted.
The power of suspension by a thread is also possessed by certain
of the Cyclostomatidae, by some Cerithidea, several Rissoa and
other marine genera, prominent among which is Litiopa bombyx,
whose name expresses its power of anchoring itself to the Sargasso
weed by a silken thread of mucus. Several species of slugs are
known to be able to let themselves down by threads from the
branches of trees. Limax arborum is especially noted for this
property, and has been observed suspended in pairs during the
breeding time. According to Binney, all the American species of
Limax, besides those of Tebennophorus, possess this singular
property. Limax arborum appears to be the only slug which has been
noticed to ascend, as well as descend, its thread. It has also been
observed[45] that when this species is gorged with food, its slime is
thin and watery, and unable to sustain its weight, but that after the
process of digestion has been performed, the mucus again becomes
thick and tenacious. It appears therefore that when the animal is
hungry and most in need of the power of making distant excursions
in search of food, its condition enables it to do so, but that when no
such necessity is pressing, the thread-forming mucus is not
secreted, or is perhaps held in suspense while the glands assist in
lubricating the food before digestion.[46]
Food of Land and Fresh-water Mollusca.—Arion ater, the great
black slug, although normally frugivorous, is unquestionably
carnivorous as well, feeding on all sorts of animal matter, whether
decaying, freshly killed, or even in a living state. It is frequently
noticed feeding on earthworms; kept in captivity, it will eat raw beef;
it does not disdain the carcases of its own dead brethren. An old
man near Berwick-on-Tweed, going out one morning to mow grass,
found a black slug devouring, as he supposed, a dead mouse. Being
of an inquisitive turn, and wishing to ascertain if it were really thus
engaged, he drew the mouse a little back. When he returned in the
evening, the mouse was reduced almost to a skeleton, and the slug
was still there.[47] Indeed it would seem almost difficult to name
anything which Arion ater will not eat. Dr. Gray mentions[48] a case
of a specimen which devoured sand recently taken from the beach,
which contained just enough animal matter to render it luminous
when trodden on in the dark; after a little time the faeces of the slug
were composed of pure sand, united together by a little mucus. A
specimen kept two days in captivity was turned out on a newspaper,
and commenced at once to devour it. The same specimen ate dead
bodies of five other species of slugs, a dead Unio, pupae of
Adimonia tanaceti, part of the abdomen of a dragon-fly, and Pears’
soap, the latter reluctantly.[49]
According to Simroth[50] and Scharff[51] the food of several of our
British slugs, e.g. Limax maximus, L. flavus, Arion subfuscus, A.
intermedius, consists of non-chlorophyllaceous substances only,
while anything containing chlorophyll is as a rule refused. On the
other hand L. agrestis and Amalia carinata feed almost entirely on
green food, and are most destructive in gardens. The latter species
lives several inches under ground during the day, and comes to the
surface only at night. It is largely responsible for the disappearance
of bulbs, to which it is extremely partial. L. marginatus (= arborum
Bouch.) feeds exclusively on lichens, and in captivity absolutely
refuses green leaves and a flesh diet. It follows therefore, if these
observations are correct, that the popular notions about slugs must
be revised, and that while we continue to exterminate from our
gardens those species which have a taste for chlorophyll, we ought
to spare, if not encourage those whose tastes lie in the opposite
direction.
Limax agrestis has been seen devouring the crushed remains of
Arion ater. Five specimens of the same species were once noticed
busily devouring a May-fly each, and this in the middle of a large
meadow, where it may be presumed there was no lack of green
food. The capture and eating of insects by Mollusca seems very
remarkable, but this story does not stand alone. Mr. T. Vernon
Wollaston once enclosed in a bottle at least three dozen specimens
of Coleoptera together with 4 Helix cantiana, 5 H. hispida, and 1 H.
virgata, together with an abundant supply of fresh leaves and grass.
About a fortnight afterwards, on the bottle being opened, it was
found that every single specimen of the Coleoptera had been
devoured by the snails.[52] Amalia marginata in captivity has been
fed upon the larvae of Euchelia jacobaeae, eating three in two hours.
[53]
Limax maximus (Fig. 19) has been seen frequently to make its
way into a dairy and feed on raw beef.[54] Individuals kept in
confinement are guilty of cannibalism. Mr. W. A. Gain kept three
specimens in a box together, and found one of them two-thirds
eaten, “the tail left clean cut off, reminding one of that portion of a
fish on a fishmonger’s stall.” That starvation did not prompt the crime
was proved by the fact that during the preceding night the slug had
been supplied with, and had eaten, a considerable quantity of its
favourite food. On two other occasions the same observer found one
of his slugs deprived of its slime and a portion of its skin, and in a
dying condition.[55] An adult L. maximus, kept for thirty-three days in
captivity with a young Arion ater, attacked it frequently, denuded it of
its slime, and gnawed numerous small pieces of skin off the body
and mantle.[56] The present writer has found no better bait for this
species on a warm summer night than the bodies of its brethren
which were slain on the night preceding; it will also devour dead
Helix aspersa. Mr. Gain considers it a very dainty feeder, preferring
fungi to all other foods, and apparently doing no harm in the garden.
Fig. 19.—Limax maximus L. PO, pulmonary
orifice: × ⅔.
Limax flavus, which is fond of inhabiting the vicinity of cellars,
makes its presence most disagreeable by attacking articles of food,
and especially by insinuating itself into vessels containing meal and
flour.[57] It is particularly partial to cream.
Slugs will sometimes bite their captor’s hands. Mr. Kew relates
that a Limax agrestis, on being stopped with the finger, while
endeavouring to escape from the attack of a large Arion, attempted
to bite fiercely, the rasping action of its radula being plainly felt.
According to the same authority, probably all the slugs will rasp the
skin of the finger, if it is held out to them, and continue to do so for a
considerable time, without however actually drawing blood.[58] While
Mr. Gain was handling a large Arion ater, it at once seized one of the
folds of skin between the fingers of the hand on which it was placed;
after the action of the radula had been allowed to continue for about
a minute, the skin was seen to be abraded.[59] Another specimen of
Arion ater, carried in the hand for a long time enclosed in a dock leaf,
began to rasp the skin. The operation was permitted until it became
too painful to bear. Examination with a lens showed the skin almost
rasped away, and the place remained tender and sore, like a slight
burn, for several days.[60]
Helix pisana, if freshly caught, and placed in a box with other
species, will set to work and devour them within twenty-four hours.
The present writer has noticed it, in this position, attack and kill large
specimens of H. ericetorum, cleaning them completely out, and
inserting its elongated body into the top whorls of its unfortunate
victims in a most remarkable manner. Amongst a large number of
species bred in captivity by Miss F. M. Hele,[61] was Hyalinia
Draparnaldi. In the first summer the young offspring were fed on
cabbage, coltsfoot, and broadleafed docks. They would not
hibernate even in the severest frosts, and, no outdoor food being
available, were fed on chopped beef. This, Miss Hele thinks, must
have degenerated their appetites, for in the following spring and
summer they constantly devoured each other.
Zonites algirus feeds on decayed fruit and vegetables, and on
stinking flesh.[62] Achatina panthera has been known to eat meat,
other snails (when dead), vegetables, and paper.[63] The common
Stenogyra decollata of the South of Europe has a very bad character
for flesh-eating habits, when kept in captivity. Mr. Binney[64] kept a
number for a long time as scavengers, to clean the shells of other
snails. As soon as a living Helix was placed in a box with them, one
would attack it, introduce itself into the upper whorls, and completely
remove the animal. One day a number of Succinea ovalis were left
with them for a short time, and disappeared entirely! The Stenogyra
had eaten shell as well as animal. This view of Stenogyra is quite
confirmed by Miss Hele, who has bred them in thousands. “I can
keep,” she writes,[65] “no small Helix or Bulimus with them, for they
at once kill and eat them. They will also eat raw meat.”
Even the common Limnaea stagnalis, which is usually regarded
as strictly herbivorous, will sometimes betake itself, apparently by
preference, to a diet of flesh. Karl Semper frequently observed the
Limnaeae in his aquarium suddenly attack healthy living specimens
of the common large water newt (Triton taeniatus), overcome them,
and devour them, although there was plenty of their favourite
vegetable food growing within easy reach.[66] The same species has
also been noticed to devour its own ova, and the larvae of Dytiscus.

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Hazzard's Geriatric Medicine and

Gerontology, 8th ed 8th Edition Kevin

Jeffrey B. Halter • Joseph G. Ouslander • Stephanie Studenski

University of Pittsburgh School of Medicine Editor Emeritus and Senior Advisor

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Principles of Palliative Medicine and Ethics

Organ Systems and Diseases

Fig. 13.—Illustrating the development of the fresh-water genus

Origin of the Land Fauna

Fig. 16.—Two rows of the radula of Littorina littorea L., × 72.

Fig. 17.—Two rows of the radula of Cyclophorus sp., India, ×

The majority of the Land Mollusca are probably more sensitive

Endurance of Heat and Cold.—The Mollusca are capable, at

Vitrina is one of our most hardy molluscs, and may be observed

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