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Reptile Biodiversity Standard Methods for Inventory and
Monitoring 1st Edition Dr. Roy W. Mcdiarmid (Editor)
Digital Instant Download
Author(s): Dr. Roy W. McDiarmid (editor), Dr. Mercedes S. Foster (editor),
Dr. Craig Guyer (editor), Dr. J. Whitfield Gibbons (editor), Dr. Neil Chernoff
(editor)
ISBN(s): 0520266714
Edition: 1
File Details: PDF, 46.23 MB
Year: 2012
Language: english
 Reptile Biodiversity
STANDARD METHODS FOR INVENTORY AND MONITORING

Edited by

ROY W. MCDIARMID
MERCEDES S. FOSTER
CRAIG GUYER
J. WHITFIELD GIBBONS
NEIL CHERNOFF

U NIVER SIT Y OF CALIFOR NIA PR ESS

Berkeley Los Angeles London
R EP TI LE B IO D IV ERS IT Y
 Reptile Biodiversity
STANDARD METHODS FOR INVENTORY AND MONITORING

Edited by

ROY W. MCDIARMID
MERCEDES S. FOSTER
CRAIG GUYER
J. WHITFIELD GIBBONS
NEIL CHERNOFF

U NIVER SIT Y OF CALIFOR NIA PR ESS

Berkeley Los Angeles London
University of California Press, one of the most distinguished Library of Congress Cataloging-in-Publication Data
university presses in the United States, enriches lives around the
Reptile biodiversity : standard methods for inventory and
world by advancing scholarship in the humanities, social sciences,
monitoring / edited by Roy W. McDiarmid . . . [et al.].
and natural sciences. Its activities are supported by the UC Press
p. cm.
Foundation and by philanthropic contributions from individuals
Includes bibliographical references and index.
and institutions. For more information, visit www.ucpress.edu.
ISBN 978- 0-520-26671-1 (cloth : alk. paper)
University of California Press 1. Reptile populations– Research–Methodology. 2. Reptiles–
Berkeley and Los Angeles, California Conservation. 3. Animal diversity conservation. I. McDiarmid,
Roy W.
University of California Press, Ltd.
QL645.6.R47 2012
London, England
597.9'072–dc22 2011011205
© 2012 by the Regents of the University of California
19 18 17 16 15 14 13 12
10 9 8 7 6 5 4 3 2 1

The paper used in this publication meets the minimum requirements

of ANSI/NISO Z39.48-1992 (R 1997) (Permanence of Paper).

Cover photographs (clockwise from left to right): Diploglossus

bilobatus (Tortuguero, Costa Rica), Terrapene carolina (Savannah River
Site, South Carolina), Leptophis ahaetulla (Tortuguero, Costa Rica),
photos by John D. Willson; Corallus hortulanus (Cerro de la Neblina,
Venezuela), photo by Roy W. McDiarmid; Pseustes sulfureus (Manu
National Park, Peru), photo by Fiona A. Wilkinson, © Maquina
Images.
 CONTENTS

AUTHORS AND CONTRIBUTORS vii PART THREE

FOREWORD ix Sampling Reptile Diversity
RICK SHINE
11 Techniques for Reptiles in Difficult-to- Sample
PREFACE xi Habitats / 167
ROBERT E. LOVICH, Chapter Coordinator
PART ONE
12 Statistical Properties of Techniques and
Introduction Validation / 197
1 Studying Reptile Diversity / 3 GORDON H. RODDA
MERCEDES S. FOSTER, ROY W. MCDIARMID,
AND NEIL CHERNOFF 13 Standard Techniques for Inventory and
Monitoring / 205
2 Reptile Diversity and Natural History: MERCEDES S. FOSTER, Chapter Coordinator
An Overview / 7
ROY W. MCDIARMID 14 Parametric Analysis of Reptile Biodiversity
Data / 273
CHAD L. CROSS, NATALIA ANANJEVA, NIKOLAI L. ORLOV,
PART TWO
AND ANTONIO W. SALAS
Planning a Diversity Study
3 Study Design and Sampling / 27 15 Population Size and Demographics / 283
ROBERT N. FISHER AND MERCEDES S. FOSTER, GORDON H. RODDA

Chapter Coordinators
16 Monitoring Exploited Species / 323
4 Dealing with Associated Data / 51 LEE A. FITZGERALD
MERCEDES S. FOSTER AND ROBERT N. FISHER,
Chapter Coordinators
PART FOUR
5 Finding and Capturing Reptiles / 77 Conclusions
LEE A. FITZGERALD
17 Reptile Biodiversity: Where Do We Go from
6 Voucher Specimens / 89 Here? / 335
ROBERT P. REYNOLDS AND ROY W. MCDIARMID ROY W. MCDIARMID AND MERCEDES S. FOSTER

7 Preparing Reptiles as Voucher Specimens / 95 APPENDIX I Selected Institutions with Significant

MERCEDES S. FOSTER, Chapter Coordinator
Collections of Reptiles 341
8 Dealing with Live Reptiles / 127 MERCEDES S. FOSTER AND ROY W. MCDIARMID
MERCEDES S. FOSTER, Chapter Coordinator
APPENDIX II Websites of Interest 345
9 Marking Reptiles / 143 MERCEDES S. FOSTER
MICHAEL V. PLUMMER AND JOHN W. FERNER

10 Determining Age, Sex, and Reproductive LITER ATURE CITED 349

Condition / 151 ADDRESSES OF AUTHORS AND CONTRIBUTORS 391
ROBERT N. REED AND ANTON D. TUCKER INDICES 395

v
This page intentionally left blank
 AUTHORS AND CONTRIBUTORS

THOMAS S. B. AKRE Longwood University HAROLD F. HEATWOLE North Carolina State University
NATALIA ANANJEVA Zoological Institute, Russian Academy ROBERT D. KENNEY University of Rhode Island
of Sciences F. WAYNE KING Florida Museum of Natural History
ANDREA ATKINSON National Park Ser vice JOHN LEE The Academy at Roosevelt Center
RALPH W. AXTELL Southern Illinois University ROBERT E. LOVICH Naval Facilities Engineering Command
SEAN J. BARRY University of California - Davis WILLIAM E. MAGNUSSON Instituto Nacional de Pesquisas da
AARON M. BAUER Villanova University Amazônia
STEVEN J. BEAUPRE University of Arkansas MARCIO MARTINS Universidade de São Paulo
CHRISTOPHER BROWN USGS Western Ecological Research FRANK J. MAZZOTTI University of Florida
Center ROY W. MCDIARMID USGS Patuxent Wildlife Research Center
GORDON M. BURGHARDT University of Tennessee - JOSEPH C. MITCHELL Mitchell Ecological Research Ser vice
Knoxville
MILAN MITROVICH USGS Western Ecological Research Center
NEIL CHERNOFF Environmental Protection Agency
JOHN C. MURPHY Field Museum
CHAD L. CROSS University of Nevada - Las Vegas
HENRY MUSHINSKY University of South Florida
INDRANEIL DAS Universiti Malaysia Sarawak
CRISTIANO NOGUEIRA Universidade de Brasilia
DALE DENARDO Arizona State University
NIKOLAI L. ORLOV Zoological Institute, Russian Academy
JAMES R. DIXON Texas A&M University (emeritus) of Sciences
MAUREEN A. DONNELLY Florida International University CHARLES R. PETERSON Idaho State University
MICHAEL E. DORCAS Davidson College MICHAEL V. PLUMMER Harding University
Thomas More College and Cincinnati
JOHN W. FERNER CHRISTOPHER J. RAXWORTHY American Museum of Natural
Museum of Natural History History
ROBERT N. FISHER USGS Western Ecological Research Center ROBERT N. REED USGS Fort Collins Science Center
LEE A. FITZGERALD Texas A & M University ROBERT P. REYNOLDS USGS Patuxent Wildlife Research Center
MERCEDES S. FOSTER USGS Patuxent Wildlife Research Center CARLTON J. ROCHESTER USGS Western Ecological Research
JACK FRAZIER Smithsonian Institution Center
SCOTT L. GARDNER University of Nebraska GORDON H. RODDA USGS Fort Collins Science Center
J. WHITFIELD GIBBONS University of Georgia D. CRAIG RUDOLPH U. S. Forest Ser vice
J. STEVE GODLEY Cardno ENTRIX, Inc. ANTONIO W. SALAS Center for Conservation Biology– Peru
KIM GRAY- LOVICH Zoological Society of San Diego ALAN H. SAVITZKY Old Dominion University
HARRY W. GREENE Cornell University JAMES A. SCHULTE II Clarkson University
PATRICK T. GREGORY University of Victoria RICK SHINE University of Sydney
CRAIG GUYER Auburn University C. ROBERT SHOOP Cumberland Island Museum (deceased)
LEE-ANN C. HAYEK Smithsonian Institution LORA L. SMITH Joseph W. Jones Ecological Research Center
WILLIAM K. HAYES Loma Linda University DAVID A. STEEN Auburn University

vii
BRIAN K. SULLIVAN Arizona State University JOHN D. WILLSON University of Georgia
ANTON D. TUCKER Mote Marine Laboratory THOMAS P. WILSON University of Tennessee - Chattanooga
WILLIAM A. VELHAGEN, JR. New York University JAMES H. YANTIS Texas Parks & Wildlife Department (retired)
RICHARD C. VOGT Instituto Nacional de Pesquisas da Amazônia CAMERON A. YOUNG Center for Snake Conservation
HAROLD K. VORIS Field Museum ROBERT T. ZAPPALORTI Herpetological Associates, Inc.

viii AUTHORS AND CON TRIBUTORS

 FOREWORD

RICK SHINE

This is the book that I desperately needed at the beginning of turtles, crocodiles, or tuataras for a very long time, and these
my scientific career. Like most other young herpetologists, I independent evolutionary histories have produced animals
had a pretty simple set of ideas about how to gather data— I’d that differ from each other in many ways. Even within each
just go out there, look for snakes, find some, catch them, and of the major lineages, reptiles are amazingly diverse in almost
then write down anything that seemed useful (such as their every aspect. One inevitable result is that an equally diverse
sex or body size). And somehow or other, once I’d been doing array of methods is needed for studies of those animals—for
that for long enough, I’d have a data set that could then tell example, the techniques that enable one to most effectively
all kinds of interesting stories about the biology of the crea- find, collect, and study a blind snake embryo differ in sub-
tures in question. Fortunately, there were wiser heads around— stantial ways from the techniques needed to investigate adult
notably my doctoral supervisor, Hal Heatwole, who not only saltwater crocodiles. That diversity creates an enormous chal-
is still an active researcher but also has contributed to this lenge for anyone who attempts a book like the current vol-
volume. And after talking with those more experienced her- ume. Fortunately, the critical importance of the task has en-
petologists, it dawned on me that I actually needed to think abled the editors to fi nd a stellar list of contributors, including
about what I was going to do, and how I might ultimately use many of the world’s most experienced field herpetologists.
the data, before I started my fieldwork. The task is urgent. In many parts of the world, reptile pop-
One of the glories of fieldwork is serendipity: you never re- ulations are being menaced by a cocktail of threatening pro-
ally know what you might discover (famously, Albert Einstein cesses, ultimately brought about by human activities. Reptiles
commented that if we knew what we were doing, it wouldn’t are not unique in their vulnerability to threats such as habi-
be research). My own career has been full of such surprises: tat degradation, climate change, and invasive species— but
I’ve been astonished by some peculiar facet or other in the bi- the central role of reptiles in many ecosystems is rarely ap-
ology of almost every species that I have studied. At first sight, preciated. Popular attention tends to focus on endothermic
a standardizing of methods— one of the main points of this vertebrates, especially large mammals and brightly colored
book—might seem counter to that spirit of free inquiry. It’s birds. If we look to the tropics, however—the parts of the
not. If your data set is gathered with a robust sampling design world that are the most biodiverse and that are subject to the
and in a way that can be compared directly to other data sets most intense threats— the diversity and abundance of ecto-
gathered at other places and other times on the same or differ- thermic (“cold-blooded”) vertebrates like reptiles and amphib-
ent species, the opportunities to discover something genuinely ians vastly exceeds the numbers and types of birds and mam-
new and exciting are magnified a thousand-fold. And it is only mals. Our current estimates of reptile biodiversity in most
by bringing separately gathered data sets together that we can tropical regions also are far too low, with new species being
address many issues that are critical for conservation and man- described every month. The small body sizes, low metabolic
agement. For example, some of the most important challenges rates, and sedentary behaviors of many reptile species have
to the viability of reptile populations— such as climate change resulted in many taxa being restricted to very small areas,
or the overall impact of invasive species or diseases— operate at some of which undoubtedly have been destroyed long before
timescales and spatial scales too large for any single field study. the species within them were ever seen by scientists. As ecto-
Field-based surveys on any organisms need robust sam- therms, reptiles rely upon ambient temperature regimes for
pling designs as well as methods that are logistically feasible many aspects of their day-to-day lives, and climate change may
and ethically acceptable. But the need is greater— and the pose a massive challenge to the continued existence of many
challenge greater—for reptiles than for many other types of species.
animals. As any phylogenetic purist will tell you, there’s no Reptilian ectothermy also poses a huge challenge to any
such thing as a “reptile” in evolutionary terms— squamates attempt to establish and describe standard techniques for
(lizards and snakes) haven’t shared a common ancestor with biodiversity monitoring. The intimate dependence of reptile

ix
behavior on local weather conditions means that surveys con- sible to apply standard techniques for biodiversity monitoring,
ducted at different times can yield massively different num- but they do place a premium on refining methods in light of
bers of animals, and the ability to remain inactive and unfed local conditions and ecological circumstances.
for weeks or months at a time makes these animals a night- In summary, then, monitoring reptile populations in stan-
mare to survey quantitatively— a high proportion of the pop- dardized, repeatable ways is not an easy task—but it’s a criti-
ulation may be virtually inaccessible to an observer for most cally important one. Unless we can accurately measure what’s
of the year. The reptiles of severely seasonal climatic regimes out there in the wild, we cannot detect how, when, where, and
offer an obvious example: calculating abundances of garter why things are going wrong. There will never be a simple
snakes on the Manitoba prairies is impossible for the 8 months “one size fits all” formula for asking questions of nature, but
per year that they spend underground, before emerging during standardization—and utilization of the most powerful sam-
the brief warm period. But many examples are less obvious: for pling designs— can add enormously to the value of the infor-
example, pit-vipers on the small island of Shedao in northern mation that is collected. This book, encapsulating the hard-
China are active only for a few weeks each year in spring and won experience of expert field herpetologists, offers many ideas
autumn, because these are the only times that their main and protocols that will improve the quality of our science and
prey—migrating passerine birds—pass through the island in ultimately help us to conserve the organisms that we cherish.
seasonal migrations. Any attempt to monitor abundances or
population attributes in such systems must incorporate knowl- Rick Shine
edge of their ecology, and hence of the appropriate periods for University of Sydney
sampling. Species’ peculiarities like this don’t make it impos- May 2010

x FOREWORD
 P R E FAC E

Historical Perspective protocols in a single volume in a user-friendly format would

save time and funds in the planning of any inventory or
This volume is the fourth in a series of handbooks developed monitoring project because it would not be necessary for in-
to identify and present standard methods for the inventory vestigators to “reinvent the wheel.” In addition, a user-friendly
and monitoring of biological diversity. Ground-level informa- handbook would be readily available for training field practi-
tion about biodiversity provides the basis for informed deci- tioners, technicians, and students.
sions about the protection, sustainable use, or development of With the broad participation and cooperation of the com-
specific areas or species and for the rehabilitation or restora- munity of amphibian biologists, the fi rst volume in the series
tion of degraded environments. Yet our knowledge of biodi- (Heyer et al. 1994a), focusing on Amphibians, was produced.
versity, including the geographical and habitat distributions Duplicating the model, volumes on standardized methods for
of most organisms, ranges from imprecise and incomplete to use with mammals and fungi followed. The present volume
nonexistent, and many habitats and biotas are disappearing deals with reptiles, a group that represents a challenge for
or being degraded faster than their biodiversity can be inven- many reasons, including the highly disparate morphologies,
toried. Funds for such work tend to be scarce, and the number ecologies, and behaviors found among extant groups and
of trained scientists, technicians, and managers is limited, the scarcity of systematic, quantitative baseline data against
particularly in the least- explored areas of developing coun- which to measure population changes that may have occurred
tries. By collecting and recording data according to standard over time. Yet reptile populations face a multitude of threats,
methods and protocols and analyzing them in standard ways, including habitat destruction, overcollecting for the pet trade,
the utility of the data that are being gathered is maximized. excess and often unregulated commercial harvest for food,
Data from diverse studies can be compared across sites and skins, or trinkets, incidental anthropogenic mortality, and gen-
through time; they can also be combined to provide regional eral animosity toward some groups (e.g., poisonous snakes) that
or even larger-scale coverage. Given the added value of stan- leads to their wanton killing. Consequently, when Neil Cher-
dardized data, it is surprising that no generally accepted and noff approached Mercedes Foster and Roy McDiarmid about
widely used standard methods for sampling the diversity of the possibility of producing a handbook dealing with meth-
most organisms are available. ods for reptiles, their responses were extremely enthusiastic.
This lack was particularly evident when suspicions of global Foster convened a core committee (Roy McDiarmid, Craig
declines in amphibian populations arose in the late 1980s. Guyer, and Neil Chernoff) to develop a preliminary plan for
Verification of supposed population changes was hampered the book and seek comments and expressions of interest from
by the absence of quantitative baseline population data against the greater community of reptile biologists. The committee
which to compare current population levels. In a (belated) at- subsequently invited individuals who had experience with
tempt to fill that void, Mercedes Foster initiated the Biodiver- the inventory and monitoring of reptiles to attend a work-
sity Methods Handbooks project in 1990. She assembled a shop to complete the outline for the book and assume re-
group of amphibian biologists and encouraged them to pro- sponsibility for preparing chapters. The workshop was held
duce a volume identifying, detailing the use of, and advocating from 29 to 31 August 2002 in Arlington, Virginia. Partici-
a series of standardized methods for inventorying and monitor- pants included Neil Chernoff, Robert Fisher, Lee Fitzgerald,
ing the biodiversity of amphibians. The anticipated book would Mercedes Foster, Whit Gibbons, Craig Guyer, Lee-Ann Hayek,
cover biodiversity sampling in all habitats and ecosystems John Lee, Robert Lovitch, Roy McDiarmid (Workshop Chair),
wherever amphibians might occur throughout the world. It Joe Mitchell, Jim Nichols, Charles Peterson, Christopher Rax-
would also include sections on project design, data analysis, worthy, Robert Reed, and Gordon Rodda. At that time, McDi-
data and specimen standards, GIS, and similar topics gener- armid, Foster, Guyer, Gibbons, and Chernoff assumed re-
ally applicable to all biodiversity studies. Having the standard sponsibility for compiling and editing the publication.

xi
 Despite the best of intentions, completion of the manu- obtain copies of publications. Fiona Wilkinson (USGS/PWRC)
script was delayed because of illness, lack of funds, and author helped with the verification of literature references and sup-
delays. It was fi nally sent to the Press on 19 December 2010. plier contact information, prepared several of the figures, and
Ultimately, 70 reptile biologists and others with overwhelm- assisted Foster with a myriad of miscellaneous tasks related to
ing collective expertise and from six countries contributed to preparation of the manuscript. James D. Nichols (USGS/
the volume. The participation of so many highly experienced PWRC) and Sally T. Murphy contributed in other ways.
individuals helped to ensure that the book provides compre- We are deeply indebted to the authors for contributing the
hensive and up-to- date coverage of sampling methodologies manuscripts that make up this volume and greatly appreciate
and data analyses, and we believe that the volume will make their willingness to participate in the project. We are espe-
a major contribution to the study of reptile biodiversity. How- cially grateful to the participants who submitted their manu-
ever, sampling methods and analytical procedures are never scripts on time and maintained their equanimity in the face
static. We encourage investigators interested in reptilian bio- of delays caused by those who did not. Finally, we would like
diversity to refine and improve the methods covered in this to thank Chuck Crumly, our Editor at the University of Califor-
book, and we look forward to the development of new and nia Press, for his infinite patience during the preparation of this
innovative procedures. volume and his faith in our ability to finish the manuscript. We
also thank Lynn Meinhardt and Francisco Reinking, our Edito-
rial Coordinator and Project Editor at the Press, respectively,
Acknowledgments and Edward Wade, Production Editor at Westchester Book Ser-
vices, all of whom answered our questions and provided ex-
Many people worked tirelessly to bring this ambitious project tremely helpful advice and direction.
to completion, and we offer them our heartfelt thanks. We are The Environmental Protection Agency provided generous
especially grateful to Paul Ustach for providing the delightful financial support during the early stages of this project, in-
cartoons that liven up the volume. We also thank all those field cluding funding for the workshop. Other financial and admin-
herpetologists who generously allowed us to use their photo- istrative support was provided by the USGS Patuxent Wildlife
graphs and authors and publishers who provided permission to Research Center and the Smithsonian Institution’s National
reprint previously published material. Museum of Natural History. Any royalties from the book will
Lynda J. Garrett of the USGS Patuxent Wildlife Research be donated to Partners in Amphibian and Reptile Conservation
Center (USGS/PWRC) Library worked miracles in locating (PARC) to support its national and international activities
and obtaining copies of reports from the gray literature, Fed- (http://www.parcplace.org/about.html).
eral and State documents, and mainstream publications. Pa-
tricia L. Lasker, Martha A. Rosen, and David T. Steere, Jr., of The Editors
the Smithsonian Institution’s National Museum of Natural Washington, D. C.
History Branch Library also gave generously of their time to December 19, 2010

xii PREFACE
 PART ONE

INTRODUCTION
This page intentionally left blank
 CHAPTER ONE

Studying Reptile Diversity

MERCEDES S. FOSTER, ROY W. M C DIARMID, and NEIL CHERNOFF

Biological diversity is the key to the maintenance of the world as

we know it.
E. O. WILSON, 1992

Sentient species that we are, Homo sapiens is able to understand

the shape, extent, and value of Earth’s biodiversity. Indeed, we
have a responsibility, as well as a self-interest, to value it.
RICHARD E. LEAKEY and ROGER LEWIN, 1995

Setting the Stage / 3 they hoped might be achieved by 2010, the United Nations–
Importance of Standardization / 4 designated International Year of Biodiversity. They recognized,
Intended Audience / 5 however, that in most instances the available taxonomic infor-
mation was too limited to form a basis for decisions about the
conservation and sustainable use of our living resources. To
Setting the Stage help reduce this “taxonomic impediment,” they established
the Global Taxonomy Initiative to train taxonomists, expand
This volume is the fourth in a series of publications dealing scientific collecting, and increase the pace of taxonomic work,
with standard methods for inventorying and monitoring the particularly the description of new species (SCBD 2008).
biodiversity of different taxa (e.g., see Heyer et al. 1994a; Wilson After the Rio meeting, many countries rose to the challenge,
et al. 1996; Mueller et al. 2004). The impetus for the fi rst vol- designating new protected areas, developing environmental
ume was the sudden realization in the late 1980s and early education programs, and increasing efforts to manage exploited
1990s that amphibian populations were declining globally, and threatened ecosystems. In addition, countries, companies,
for unknown reasons. Determining the scope (numbers of spe- and conservationists began placing more emphasis on sustain-
cies affected) and the magnitude of the declines was hampered able development and habitat restoration rather than just on
by the absence of baseline data with which to compare con- the protection of pristine areas, but at the same time recogniz-
temporary population levels and by the lack of comparability ing the importance of ecosystem ser vices (carbon sequestra-
among the data sets that were available. tion, clean air and water, soil maintenance, pollinator popula-
About the time that the amphibian declines were first being tions and food production, sources of raw materials with newly
recognized, there were dramatic surges in interest and concern recognized uses) whose economic value has been largely unac-
about the worldwide loss of biodiversity, even though various knowledged. Nevertheless, by the time you read this the 2010
individuals and institutions had been sounding the alarm for deadline will have passed, and although rates have perhaps
decades. In 1986, the National Academy of Sciences and the slowed, habitats continue to shrink, species continue to de-
Smithsonian Institution sponsored the National Forum on cline or go extinct (Hoffman et al. 2010; Marton-Lefèvre 2010),
BioDiversity, held in Washington, DC, on September 21–24, and government and conservation groups continue to base
and published the proceedings of that forum (see Wilson 1988). decisions regarding conservation, management, and develop-
In 1992 (3–14 June), the United Nations held the Conference ment on woefully inadequate and incomplete data about the
on Environment and Development (UNCED) in Rio de Ja- distributions and population statuses of species. At a time
neiro, Brazil. The attendees, who numbered between 20,000 when our need to use our resources more efficiently and main-
and 30,000, represented the governments of 178 countries as tain healthy populations of the species and ecosystems of great-
well as nongovernmental organizations (NGOs) and the news est value to the health of the planet, we still do not know the
media. One product of that meeting was The United Nations species composition of most ecosystems, understand the func-
Convention on Biological Diversity ( http://www.cbd.int/con tion of each in those ecosystems and how they maintain it, or
vention/text/ ), often referred to as the Biodiversity Treaty. appreciate the impact the loss of any one of them might have.
Recurring themes of the conference and the Convention were Without such information, conserving biodiversity, restoring
that countries needed to conserve their biological diversity
and use their biological resources in a sustainable manner. At a
Reptile Biodiversity: Standard Methods for Inventory and Monitoring, edited by
meeting in April 2002, the Conference of the Parties (the Roy W. McDiarmid, Mercedes S. Foster, Craig Guyer, J. Whitfield Gibbons,
decision-making body of the Convention) set as a goal a sig- and Neil Chernoff. Copyright © 2012 by The Regents of the University of
nificant reduction in the rate of loss of biodiversity, which California. All rights of reproduction in any form reserved.

3
degraded habitats, and reconstituting ecosystems will be et al. 2002; Carreira et al. 2005; Das 2010). We hope that this
impossible. volume will contribute to that effort.
Some 25 years after recognizing the global decline of am- In this volume we provide information on carry ing out
phibian populations, biologists and conservationists are be- standardized quantitative surveys of reptile biodiversity that
coming increasingly aware of declines in reptile populations can be compared across habitats and through time. We include
(Gibbons et al. 2000; Yardley 2007; Sinervo 2010). And once chapters addressing issues to consider when planning a study,
again, we are faced with an absence of baseline data with selecting sampling techniques, performing a survey, and ana-
which to make comparisons. Nearly 9,100 species of reptiles lyzing the data obtained. It is our fervent hope that this volume
are known, and new taxa continue to be described at a prodi- will motivate scientists, managers, conservationists, and nature
gious rate. Uetz (2010) estimated that on average nearly 100 lovers, whether employed by governments or conservation
new species of reptiles were described each year between 1998 groups or simply aficionados of reptiles, to carry out surveys of
and 2008, a rate higher than that for any 10-year interval since the biodiversity of this poorly know but exceedingly impor-
1758, and it continues to accelerate. Reptiles make up more tant group.
than half of the biomass in some ecosystems and must play a
significant role in ecosystem function. Yet, we have no knowl-
edge of many components of their basic natural history. The Importance of Standardization
lack of data may in part be attributable to the less charismatic
nature (e.g., compared to birds and mammals) of these organ- Throughout this book we advocate the use of standard
isms that may even engender fear and hostility on the part of methods for inventory and monitoring. Standard methods
some—there is no “Kermit” equivalent to speak for them. increase the comparability of data and decrease the effects of
The herpetofaunas of many areas of Europe and much of variables extraneous to a study. Nevertheless, because reptile
eastern North America have been studied for more than 150 species exhibit such a wide range of morphologies, behaviors,
years. Regional summaries published from the late 19th to and ecologies, it is impossible to identify a single technique
mid-20th centuries were followed by field guides to the species that will sample all members of a reptile community equally.
of most temperate regions. The guides included drawings of Species differ, including closely related forms, and so do groups
the species accompanied by descriptions and geographic dis- within species (e.g., males and females, juveniles and adults,
tributions. Until quite recently there have not been field guides and reproductive and nonreproductive individuals). How
or regional summaries to reptiles in most tropical regions, al- many and which techniques will be employed in a study will
though species descriptions are available and distributions depend largely on its intent and whether it focuses on a single
are moderately well known. The basic components of their species, members of a particular behavioral or ecological guild,
ecologies, in contrast, are not. For example, in a recent compre- or all members of a community. And although a single species
hensive and well-illustrated field guide to the reptiles of South- may be sampled with more than one technique, studies of
east Asia, Das (2010) included in each species account a section whole communities will require the use of several. Obviously,
titled “Habitat and Behavior.” More frequently than not, these techniques appropriate for aquatic turtles will be less than rel-
sections refer to diet, reproductive behavior, and other natural evant for sampling arboreal lizards. In fact, we encourage the
history traits with notations such as “unknown,” “nothing use of more than one technique when time, personnel, and
known,” and “unstudied.” A review of the accounts revealed other resources permit.
that 288 (67%) of 428 species of lizards in 11 families were data It is possible to sample many groups of reptiles quite appro-
deficient, with values ranging from 0 percent for a few small, priately with several different techniques or variations of a
well-studied families (e.g., Lanthanotidae and Varanidae) to single technique. And most field biologists have favorites de-
highs of 84 and 93 percent, respectively, for species in the veloped by trial and error in the field or learned at their men-
poorly know families Anguidae and Dibamidae. Similarly, 260 tor’s knee. We are not suggesting that the techniques we have
(54%) of 482 species of snakes in 12 families were data defi- identified are the only appropriate ones or even the very best
cient, ranging from 0 percent for snakes in the families Py- for a par tic ular group. Rather, we have selected those that we
thonidae and Xenopeltidae to 59 and 82 percent for species believe can best be standardized, are easiest to use with the
in the families Colubridae and Typhlopidae, respectively. Not least likelihood of error, provide the greatest return for the
surprisingly, venomous species are moderately well studied, effort and resources expended, and yet produce scientifically
with only about 40 percent of the species in both the Elapidae sound and statistically rigorous data. However, when com-
and the Viperidae being data-deficient. Values for lizards and paring different sites or different times, even employing a
snakes throughout the Neotropics are likely similar (RWM, standard sampling technique is insufficient if investigators
pers. observ.). Thus, while field guides are very useful for intro- do not adhere strictly to the indicated protocol, something
ducing the reptiles of an area to both professionals and ama- that we strongly encourage. We also recommend that investi-
teurs, the existence of such books in no way signals that knowl- gators record detailed metadata (i.e., data about the data), in-
edge of the animals’ biologies is even adequate. That belief, the cluding not only when and how the primary data were col-
“field guide syndrome,” rarely reflects reality. Even in parts of lected but also the locations of study sites and when and how
the world where reptiles have been well studied, basic knowl- to access them; the distribution and nature of habitats and
edge of the natural history of most species is poorly known, other environmental features within those sites that are impor-
and knowledge gaps are rarely reported in field guides. Given tant for reptiles (maps and photographs are very useful); data
the increasing rates of discovery of new taxa, the better sam- on weather conditions at the time of collecting; descriptions
pling of poorly known geographic areas, and the increasing of training provided to technicians, assistants, and volunteers;
number of local herpetologists carry ing out quality research and lists of vouchers (specimens, photographs, videos, etc.)
on reptiles in Asia, Africa, and the Americas, progress is being collected and where they are deposited. Such information sig-
made. Recent publications reflect those advances (e.g., Spawls nificantly increases the value of the primary data and enables

4 IN TRODUCTION
 Frank and Ernest used with the permission of the Thaves and the Cartoonist Group. All rights
reserved.

future investigators to duplicate previous work as closely as erature so that individuals working in especially difficult areas
possible. or habitats or with especially problematic data can access spe-
Various techniques do exist that allow investigators to ma- cialized information applicable to unusual situations. In keep-
nipulate or transform data so that it may be possible to com- ing with our theme of standardization, we have attempted to
pare them with data collected in other ways. Nevertheless, we adhere to the taxonomic names (scientific and standard En-
recommend that investigators use standard protocols when- glish) found in Crother (2008, as updated 31 March 2011),
ever possible. Our ignorance of reptile biodiversity (and the ITIS (Integrated Taxonomic Information System, http://www
general biodiversity of the planet) is enormous, and habitat .itis.gov/), and The Reptile Database (Uetz et al. 2011). Our
destruction and species loss are proceeding apace. Time is purpose is not to endorse any par tic ular naming system or
short, personnel are few, and resources are limited. The use of taxonomic decision, but rather, to minimize confusion. We
nonstandard methods to assess the biodiversity of reptiles is a have also abbreviated or eliminated sections on sources of
luxury we can no longer afford. supplies, chemical formulae, and so forth, which may have
been more extensive in other volumes, because such informa-
tion is easily available on the increasingly ubiquitous Inter-
Intended Audience net. We emphasize quantitative methods over the simple list-
ing of species present in an area because such data are of
This volume is intended for use by government personnel, much greater utility in recognizing and documenting changes
conservationists, managers, professional scientists, and ama- in population levels. We recognize, however, that quantitative
teurs throughout the world. It should be accessible to anyone surveys are not always possible, particularly when time, per-
with a college degree in biological or environmental science, sonnel, and other resources are limited. Nevertheless, any
but it should also be appropriate as a training manual in work- information that contributes to our knowledge of species’
shops for individuals with limited formal education. We view distributions (both geographically and ecologically) and sea-
it as a self- contained volume on the basis of which a survey of sonal activities is worthwhile, and we encourage researchers
reptile biodiversity can be accomplished. Nevertheless, we have to take advantage of opportunities to work in poorly known
provided extensive references to books and to the primary lit- areas.

STUDYING REP TILE DIVERSIT Y 5

This page intentionally left blank
 CHAPTER T WO

Reptile Diversity and Natural History: An Overview

ROY W. M C DIARMID

Introduction / 7 ecology of that region, paying particular attention to habitats

Turtles / 9 likely to be used by reptiles. To support such an effort, I have
Crocodilians / 10 listed regional references for each of the major groups of rep-
Lepidosaurs / 12 tiles at the end of its section. These publications will provide
an introduction to the herpetofaunas and facilitate compila-
tion of working lists of species likely to be present. Once a list
has been compiled, investigators should become familiar with
Introduction the ecology and general natural history of those species by
reading descriptive accounts in the literature. If possible, inves-
Reptiles represent one of the more successful evolutionary radia- tigators should also visit a museum with reptile collections
tions known, and their living representatives include a diverse from the targeted country and/or region (see Appendix I) to
and distinctive set of aquatic, terrestrial, and arboreal taxa that familiarize themselves with targeted species, especially those
occur nearly everywhere in the world. They encompass all that they have not encountered previously. Such an exercise
scaled tetrapods with an amniotic egg that lack hair and mam- will provide an understanding of the reptile diversity of the
mary glands (mammals) and feathers (birds). While some dis- area and also facilitate selection of suitable sampling methods
agreement about placement of turtles exists, for this presenta- and field gear to be used during the project. Such preparation
tion I have adopted a recently proposed classification that treats also increases the effectiveness of the sampling effort and,
turtles, crocodilians, tuataras, lizards, snakes, and the aberrant therefore, the accuracy with which the reptile diversity and
volant reptile group called birds (not treated further) as Reptiles abundance can be estimated.
(Fig. 1). To accompany the narrative, I have prepared tables that
The earliest fossil record of a reptile dates from the Pennsyl- briefly summarize the diversity, continental distributions, and
vanian period, about 300 million years ago. During the period general habitat use of the groups of living reptiles, except tua-
since then, many fossil lineages (not shown in Fig. 1) have be- taras: turtles, Table 1; crocodilians, Table 2; lizards, Table 3;
come extinct. Those that remain (exclusive of birds) are the and snakes, Table 4. The diversity counts (genera and species)
subject of this volume; they comprise about 9,088 living spe- are current through June 2010. These data were extracted from
cies currently placed in about 1,109 genera and distributed the literature, some of which is listed by geographic area at the
across five groups as follows: turtles (326 species), crocodilians end of each of the five group accounts; draft manuscripts and
(24), tuataras (2), lizards (5,519), and snakes (3,217). Among the species lists to which I had access; and colleagues with detailed
amniotes, the diversity of the reptiles, when compared to those knowledge of specific groups. Some of my diversity counts
of the living mammals (about 5,420) and birds (about 10,500), were checked against the summary data contained in the Rep-
attests to their evolutionary success. Because turtles, crocodil- tile Database (http://www.reptile-database.org/), but all esti-
ians, and lepidosaurs (tuataras, lizards including amphisbae- mates are mine.
nians, and snakes) are relatively distantly related, I have tried The habitat designations for each group (family or subfam-
to capture the highlights of the biological diversity of each ily) represent my best estimates of the primary habitats used
group in its own account. by their inclusive species and vary among tables according to
Understanding something about the diversity, distribution, their suitability for each group. Lists of species were compiled
habitat use, and natural history of each species of reptile that is
likely to occur in a particular region or at a specific site is es-
Reptile Biodiversity: Standard Methods for Inventory and Monitoring, edited by
sential for an inventory project to be successful. Therefore, I Roy W. McDiarmid, Mercedes S. Foster, Craig Guyer, J. Whitfield Gibbons,
strongly recommend that anyone unfamiliar with the herpe- and Neil Chernoff. Copyright © 2012 by The Regents of the University of
tofauna of a region to be surveyed review the literature on the California. All rights of reproduction in any form reserved.

7
Another Random Scribd Document
with Unrelated Content
 PSALM XXII. 359 torical; from ver. 16 to ver. 18, the
circumstances (which cannot be made to correspond to the
supposed condition) are liypothetical : " He sees Himself in their
midst, and witnesses their joy at His wasted form, and how after His
death they part and cast lots for His clothes." A very singular way,
assuredly, of determining the situation. One, according to it, would
need to have a very free hand, and to have a peculiar taste for
following every sudden idea. In the second part, the conversion of
the heathen is violently separated from its cause and occasion : "
The time will come when the people will again think upon Jehovah,
and turn to Him." The wdiole passage, from ver. 26 to ver. 31, will
merely show, " what a God He must be who has listened to such a
prayer, and to whom such praise will be rendered." Against this the
last verse is quite sufficient : — They shall make known His
righteousness, and that He hath done this. At the expression, " they
eat," ver. 29, there will have to be supplied, " the good things of
life," — arbitrarily (for the object to be eaten must be determined
from the preceding context), and in opposition to ver. 26. Other
attempts to set aside the actual state of the case by exposition, I
have already adverted to in my Christology. Among these we reckon
the assertion, which, after the example of Yenema, has been
frequently brought forward, that the sufferer in the Psalm is not yet
in the power of his enemies, but only threatened by them. The
passages which are brought forward for the purpose, viz., 11, 12,
20, 21, do not prove it : for the nearness of the trouble in ver 11, is
not contrasted with its presence, but with its distance ; trouble is
near to him who is in the midst of it ; the expression, " many hidls
have compassed me, etc.,^^ suits a victim which has been seized,
and, to cut off every hope of escape, has been surrounded by
ferocious enemies, for the purpose of inflicting the death-stroke ;
and the 20th and 21st verses only show, what of itself is obvious,
that tJiis stroke has not yet fallen. The 17th and 18th verses prove
the contrary : — according to them, his enemies have already
stripped the sufferer quite naked, so that his emaciation lies exposed
to his own eyes and to theirs, while they enjoy the miserable
spectacle, and divide his clothes among themselves. To refer, with
Rosenmiiller and others, the 18th verse merely to the proposal to
divide the clothes, will not do, irrespective of every other
consideration, on account of the connection with
 360 THE BOOK OF PSALMS. the 17th verse, where the
sufferer is represented as already stripped naked. Those who
propose to understand the 18th verse figuratively, appear to be at a
loss what to say in their embarrassment. The hypothesis of Jarchi,
Kimchi, and others, is much more tolerable, viz., that by the sufferer
we are to understand the people, or the pious part of the same. It
will afterwards come out that this hypothesis, and in a certain
measure, also, the one which refers the Psalm to David, has truth for
its foundation. But if we apply the Psalm to the people directly and
exclusively, we shall meet with insuperable difficulties. On the
supposition that the sufferer is the whole people, it will clearly be
necessary to understand, that by the troop of evil-doers, the dogs,
the lions, and the bulls, the heathen are especially and exclusively
meant ; for which idea the Psalm does not furnish one single particle
of evidence. The opposition everywhere, is between ivickedness and
uprightness : and it is quite arbitrary here, as in all the similar cases
which are so frequent in the Psalms, to turn a purely moral into a
national opposition. Further, if we suppose the whole people, or the
pious part of the same, to be the sufferer, how could he say he
would make known the name of the Lord among his brethren, that
he would praise Him in the midst of the congregation, that from him
would go forth His praise in the great assembly, that he would pay
his vows before them that fear Him? How could he exhort the
fearers of God, the whole seed of Jacob, the whole seed of Israel,
placing himself over against them, to praise the Lord for what had
happened to him"? How could he promise to the meek, to those who
seek the Lord, nothing more than the co-enjoyment of a salvation
which was primarily conferred on himself, and nothing more than the
strengthening of their faith from the same? The whole passage, from
22d to 26th verse, is, on that hypothesis, altogether unintelligible : it
is fatal to every view which removes the contents of the Psalm
entirely from the domain of individual application. Such views also
are contradicted by the strong prominence given throughout the
Psalm to what specially belongs to an individual person : the sufferer
speaks of his 21 other, his heart, his tongue, his skin, his hands, his
feet, etc. — a form of speech which can lose its proper application
only when well-defined marks show that the term employed is a
collective one.
 PSALM XXIT. 361 The view which has really prevailed in the
Cli]:istian Chiircli, is that which refers the Psalm directly and
exclusively to Christ. The author by no means regrets that he
adopted this view in the Christology. It was the easiest and the most
natural of those which were then before the world, to which his
attention was more immediately directed ; and he would not even
now hesitate for one moment to adopt it, were he limited to making
a choice among these, as he supposed he was, — having as yet
advanced but a little way on an independent footing into the depths
of the Old Testament. In addition to the views already mentioned,
there was still another, held by Calvin, Melancthon, Amyrald, and
others, and advocated in modern times by Stier and Umbreit, — the
typical-Messianic. David, it is maintained, according to this
hypothesis, in crying to the Lord on the ground of a particular case
of distress, transfers, elevated by the spirit of Messianic prophecy,
his own being into the extreme sufferings of the hoped-for Messiah,
and speaks as the present type of the coming Deliverer. Although
the author acknowledges that in this attempt justice is done to those
considerations which may be pleaded in favour of opposing
expositions, yet he cannot but regard it as an unsuccessful attempt
at reconciliation. Such a view of the way in whicli the Psalm was
produced, appears to him as psychologically altogether
inconceivable. How David could extend his own consciousness to
that of his offspring, is conceivable enough ; but without a
destruction of the life of the soul, we cannot conceive of an
hesitation and vacillation between one's own and another's
personality. Meantime, the direct and exclusive reference of the
Psalm to Christ, presents such difficulties, that one cannot feel
perfectly satisfied with it, but is inwardly forced to look round for
some other interpretation, which may content the exegetical
conscience. We cannot, without violence, suppose the Messiah to be
introduced speaking, without any characterization whatever of His
person,— compare, for example, our remarks on the 16th Psalm.
The Psalm, moreover, is so nearly related to a number of others,
which have the sufferings of the righteous one generally for their
subject, that it appears very difficult to break its connection with
them, and to isolate it too much. Finally, what is said, in the second
part, of the consequences of the deliverance of the sufferer, is
undoubtedly far too grand to allow of its application
 362 THE BOOK OF PSALMS. to any one Israelitish
individual, and far too personal to allow of its application directly and
exclusively to the people ; and, on the other hand, the exegetical
sense cannot reconcile itself to set aside all other realizations of the
idea, that riotMng more promotes the glory of God, that nothing
more 'powerfully tends to awaken and move the spirits of men to
serve Him, than the deliverance of stiff ering onghteousness,
whether these realizations be in the experience of individuals, or in
that of the Church at large, and to confine all to the one realization
of the idea in Christ. The mighty influence, for example, which the
almost miraculous deliverance of David from the hand of Saul must
have had in quickening the fear of God, — the events also which are
recorded in Ex. xviii. 19, " And Jethro rejoiced for all the goodness
which the Lord had done to Israel, whom He had delivered out > '
the hand of the Egyptians ; and Jethro said. Blessed be the Lord,
who hath delivered you out of the hand of the Egyptians : now know
I that the Lord is greater than all gods," — in 2 Chron. xxxii. 23, "
And many (after the Lord had glorified Himself in the deliverance of
righteous Hezekiah from his enemies) brought gifts unto the Lord to
Jerusalem, and presents to Hezekiah, king of Judah, so that he was
magnified in the sight of all nations henceforth," — and those in
Dan. iii. 28, — come so obviously within the domain of the second
part, that one can scarcely rest satisfied with any interpretation
which places them altogether out of connection with it. While all
existing interpretations are thus encumbered with serious difficulties,
Ave make our escape at once, and completely, from the region of
embarrassment and constraint, if we consider the Psalm as referring
to the ideal person of the Righteous One, — a character which is
introduced more frequently throughout the Psalms than any other,
so that nothing but ignorance can raise against this interpretation
the reproach of arbitrariness. In this interpretation, justice is done to
that truth which lies at the foundation of every one of the existing
views, while, at the same time, the difficulties Avhich stand in the
way of every one of these are avoided. On this view, the case stands
as follows : " David composed this poem for the use of the Church,
like most of his other productions, on the ground-work of his own
experience, which, in this respect, had from the beginning been so
peculiarly rich. How the righteous man in this world of sin must
suffer much ; and how the Lord, when it comes to the last extremity.
 rsALM XXII. 363 gloriously delivers him ; and how his
sufferings, through the manifestation of the Divine glory in his
deliverance and in his victory over an ungodly world, subserve the
honour of God and the sanctifying of His name, and accelerate the
approach of His kingdom — this is the theme. Every particular
righteous man might appropriate to himself the consolation of this
Psalm — might expect, in his own experience, the realization of the
hopes expressed in it, in so far as the reality in him corresponded to
the idea, — in so far as he embodied in his own person the ideal
righteous man. In like manner also might the community of the
righteous, the people of the covenant, in all public troubles, draw
from it comfort, — the confident assurance, that the extremity of
trouble must at the same time be the turning-point, and that the
seed of tears must produce a rich harvest in the way of advancing
the kingdom of God. With all this the Psalm retained, on the whole,
till the coming of Christ, the character of an unfulfilled prophecy.
According to the proportion of righteousness was the proportion of
deliverance, and of blessed results for the kingdom of God. Every
temporary fulfilment pointed forward to a perfect one yet to come.
By those in whom hope in the Messiah was in general a living one,
this could be expected only in Him. The most perfect righteousness
belongs so necessarily to the idea of the INIessiah, that it could not
be present to the mind without the most distinct recognition thereof.
XoAv, in this Psalm we find righteousness represented as necessarily
connected with the severest and deepest suffering, springing out of
the natural enmity of the ungodly world. Consequently, the inference
is clear, that the Messiah, if a righteous, must also be a suffering
one. And, further, as here we find connected sufferina; rio-
hteousness and such exalted deliverance, we infer that this salvation
in the highest and fullest sense must be the lot of him who should
be the first to realize in perfection the idea of suffering
righteousness. Lastly, as the glory of God will be in proportion to the
salvation vouchsafed, it must be in the time of the jMessiah that this
will for the first time appear in all its extent and depth, as here
described. That, according to this view, justice is done to all the
references which occur in the New Testament to our Psalm (compare
Matt, xxvii. 39, 43, 46 ; Mark xv. 34 ; John xix. 24 ; Heb. ii. 11, 12 ;
and on the passage, the Christology, page 176, etc., snd besides,
also Matt, xxviii. 10, and John xx. 17, where our
 364 THE BOOK OF TSALMS. Saviour, after His resurrection,
with a significant reference to ver. 22, calls His disciples His
brethren), is clear as day, and becomes particularly obvious, when
we direct our attention to the other quotations from the Psalms in
the history of our Saviour's sufferings. Not one of them refers to a
Psalm which is of direct and exclusive Messianic import. The 69th
Psalm, which, next to the one now under consideration, is the most
remarkable, contains features which will not apply to Christ (the
strong prominence, for example, given to the sinfuhiess of the
sufferer), and which exclude the idea that our Lord and His Apostles
have given it a direct and exclusive Messianic interpretation. Still, it
is necessary to observe, that the providence of God so ordered the
circumstances, that the inward conformity of the sufferer of our
Psalm to Christ should become outwardly manifest. The Psalm would
have been fulfilled in Christ, even although the passers-by had not
shaken the head, or the mockers quoted its very words ; even
although there had been no dividing of His garments or casting lots
upon His vestures. But the striking resemblance in these particular
circumstances must be considered as an index, pointing to the
hidden, imvard resemblance. The same object subserved by this
secret guidance of Divine providence, Christ also had in view, when
He borrowed His first exclamation on the cross from the
commencement of the Psalm, and referred in His last words to its
closing sentence ; thereby impressively intimating, that the whole
Psalm was now being fulfilled. The question may very naturally be
asked. What is it that has brought such honour to our Psalm (which
even Strauss, though without a good intention, has entitled the
programme of the crucifixion of Christ) what is it that has led to its
being exalted above so many similar Psalms by which it is
surrounded, — Psalms which celebrate the contest of the righteous
in this world of sin, and the deliverance which the Lord vouchsafes
to them, and are consequently, also, indirect prophecies of Christ ;
inasmuch as every suffering that fell to the lot of a righteous man
because of his righteousness, and every deliverance which a
righteous man obtained because of his righteousness, was
presignificant of Him ? To this question a threefold answer may be
given. First, as has been suggested by Umbreit : " Among the many
Psalms which speak of the persecutions of the righteous by their
enemies, there is not one other Psalm which so
 PSALM XXII. 365 expressively and powerfully collects
together, and concentrates in one individual figure, the accumulated
pains and tortures of the sufferers in the contest with an ungodly
world." Second, those Psalms which originally refer to one particular
individual sufferer, stand one degree more remote from direct
application to the Messiah than this one, which does not first require
a separation of the idea from the individual. In like manner, the
reference to the Messiah is less prominent in those Psalms in which
the righteous man is introduced speaking, but with a reference to his
own failings and w^eaknesses. Of these no mention whatever is
made in this Psalm. Lastly, in no Psalm are the consequences which
flow from the deliverance of the righteous man painted in such
prominent and comprehensive colours as they are here. Title. To the
chief musician — on the hind of the daion of the morning — a
choicePsabn of David. The expression, nn:^'n rh^ii hv, has been
very variously interpreted. The simple remark, however, that nya,
wherever it occurs, always signifies a hind, and that it would be
perfectly arbitrary to give it any other interpretation here, so
decidedly sets aside a whole host of expositions, that it is
unnecessary even to quote them. The intei'pretation of iri'J' is in like
manner ascertained : all expositions which do not translate it by the
daicn of the morning, must at once be thrown aside. Those who
keep by the ascertained sense of the words, are generally of opinion
that these words are either the beginning of a song, or a passage
from one, the tune of which is to be sung to this Psalm : like, " The
hind of the morning." These again are divided, as to whether the
expression must be understood as denoting literally a hind, or
(according to Gesenius in the Thes.) as a poetical phrase for the
rising sun. This last interpretation is without any analogy in the
Hebrew language ; and has a very insufficient ground to rest on in
the fact that Arabic poets designate the rising sun " roe ;" and a still
weaker support in the fact that the Talmud uses the term, " the hind
of the davm of the morning," which, however, is not original, but has
obviously flown from the passage before us. This whole exposition,
however, has this against it, that there is not one single ascertained
case, in which a poem, the tune of w'hich is to be sung to the Psalm,
is quoted in the title. Only in a case of utmost necessity, therefore,
could we come to the resolution of adopting such an interpretation.
Especially, be 
 366 THE BOOK OF PSALMS. fore adopting it, would it be
necessary for us to inA'estigate whether it be not possible to
interpret the words as designative of the subject of the Psalm. On a
close examination of similar dark and enigmatical superscriptions,
especially of such as are introduced with ?]}, it almost always
appears that they demand such an interpretation. More especially in
those Psalms of which David is the author, such a reference is one
which might a priori be expected, as David was particularly fond of
indicating, by such enigmatical superscriptions, the contents and
object of his Psalms. It cannot be denied that the hind is a very
appropriate emblem of the suffering and persecuted righteous man
who meets us in the Psalm. On the one hand, the stag, or the hind
and the roe, are frequently employed as emblematical of one
persecuted or put to death. For example, 2 Sam. i. 19, David himself
says of Jonathan, " The roe, O Israel, is slain on thy high places;" —
on which clause Michaelis makes the following remark, " comparatur
Jonathan cum caprea a venatoribus confossa :" Prov. vi. 5, " Deliver
thyself as a roe from the hand of the hunter, and as a bird from the
hand of the fowler," Isa. xiii. 14. And, on the other hand, the hind
and the roe are used as emblems of loveliness, Gen. xlix. 21 ; Prov.
V. 19 ; Song of Sol. ii. 7, 9, viii. 14 ; and by the Arabians as emblems
of innocence, especially on the persecuted. In Meidani (Freytag, Th.
1, N. 148), there occurs the proverb, " eum invadat malum, non
dorcadem," him — jwt an innocent or a righteous person : and
Ferazadak (in Freytag on the passage) says, on receiving intelligence
of the death of one of his enemies : " dico ei, cum ejus mors mihi
nunciata esset : ei non dorcadi albos in arenarum tumulo (accidat)."
There is the less reason for hesitating as to this interpretation, if we
remember that David, in other places, draws from the animal
creation emblems of the sufferers and the persecuted : 1 Sam. xxvii.
20, " The king of Israel is come out to hunt a flea, as when one doth
hunt a partridge on the mountains;" xxiv. 15, "After whom is the
king of Israel come out ? After whom doest thou pm*sue ? After a
dead dog, after a flea?" and, in the title to the 56th Psalm, " on the
dumb dove among the strangers," which bears a remarkable analogy
to the passage before us. The reasons already adduced show, that it
is at least exceedingly probable that the hind may be a figurative
expression significant of suffering innocence. And it is put beyond
doubt by the fact, that
 PSALM XXII. 367 the wicked and the persecutors in this
Psalm, to the peculiar physiognomy of which belong emblems draion
from the brute creation, are designated by the terms dogs, lions,
bulls, and buffaloes. In the title of such a Psalm, we might, a priori,
expect to find such a description of the sufferer as should
correspond to that of the persecutors, especially as no such
appellation occurs in the body of the Psalm. A special argument in
favour of this interpretation is furnished by the term ''n'i?''&?, my
sti'ength, ver. 19, — a word which occurs nowhere else in Scripture,
and which seems to have been formed by the Psalmist for the sake
of the allusion to the title. The np'^K (hind) has its name from
strength, but it lacks the substance : — a creature without strength,
it is the natural prey of dogs, lions, buffaloes. But the strength which
it has not in itself it has in the Lord, who must hasten to the help of
the weak. On every other interpretation, the reference of ni^J^ to
n?"'X, which is so manifest, remains unexplained. Finally, this
reference shows at the same time that the title came from the j)en
of the author of the Psalm, and goes far to establish the originality
generally of the titles. We are led to the same result by the manifest
connection between n^X, and the expression ""bx, vX, proper/ my
Strong One, at the very opening of the Psalm, and also by the
circumstance that the symbolical designation of the sufferer in the
title exactly corresponds to those of his enemies in the Psalm itself.
All these references are so fine and significant, that they can have
proceeded only from the author himself. Hitherto we have been
discussing only the term " hind," and have left its adjunct, " the
dawn of the morning," out of sight. The generality of those who
consider the title as indicating the contents of the Psalm, trace the
connection which the hind has with the morning dawn to its being
early hunted. But this reference is too remote to admit of its being
intended by such a short expression. The only legitimate exposition
is that which is grounded on the general figurative use of the
morning dawn. That the morning dawn is used in a figurative sense,
Ave are entitled to expect from the analogy of the hind. Now, the
common idea conveyed by the figurative use of the morning dawn,
is that of " prosperity coming after misfortunes." Hence in Isa. Iviii.
8, " Then shall thy light break forth as the morning;" 10, " Then shall
thy light rise in obscurity, and thy darkness be as the noon day :"
Isa. xlvii. 11, " There shall evil come
 368 THE BOOK OF PSALMS. upon thee, the morning
whereof thou shalt not know ;" viii. 20. Hos. vi. 3, X. 15. 2 Sam.
xxiii. 4. The exprassion will thus indicate the prosperous termination
of the sufferer's condition : the suffering righteous man to whom
salvation is imparted, — a title as suitable, as exactly corresponding
to the contents, as can well be conceived. The fact so carefully
brought forward by the Evangelists, that Christ rose at the day-
dawn, — a circumstance by no means unimportant, — points to the
expression, " of the morning." The first division of the first part
begins, in the 1st and 2d verses, with the complaining question, and
the interrogative complaint, " My God, my God, why hast Thou
forsaken me?" In grounding this complaint, it is shown first, vers. 3-
8, that God is acting towards the sufferer, whom He is giving over to
destruction, in a very different manner from the way in which He
had manifested Himself, in all time past, in the experience of His
people ; and then, vers. 9 and 10, that God is as really the God of
the sufferer as He had been theirs. To this detail the prayer is next
appended, ver. 11, that God would remove the anomaly thus
demonstrated to exist, that He would not be far from the sufferer,
that He would not forsake him. Ver. 1. Ml/ God, my God, lohy hast
Thou forsaken me, far from my deliverance, far from the wo7'ds of
my groaning ? In the first clause everything depends upon defining
the idea of forsaking. This term can here signify nothing less than an
entire and complete giving up. For the trial is completely at an end,
as soon as God reveals to the sufferer that now his sufferings shall
have an end. As soon as he can say, " Thou hast heard me," he sees
that everything is right. The trial also does not consist in temporary
suffering, considered as such — this the sufferer knows 'that he
must lay his account with — but in the supposition that he has been
given up by God altogether, and for ever. Hence therefore the cry, "
Thou hast forsaken me," does not refer to an actual fact, but rests
on a conclusion which the sufferer draws from his apparently
thoroughly desperate condition, and upon the feeling of his flesh,
which cries, that now, when there is but a " hair between him and
death," everything is utterly lost. To get free from this conclusion
and this feeling, is the work that devolves on the sufferer. After he
has honestly done his part, and taken living hold of those truths
which render the forsaking altogethea* impossible,
 PSALM XXII. VEK. 1. 369 he receives from God the only
answer which can be given to his complaint, " Why hast Thou
forsaken me ?" "I have not forsaken thee, notwithstanding
appearances and feeling.'' From this exposition, it is evident that
these words, so far from being expressive of despair, are rather
destined to comiteract despair, to tear it up by the roots, when it is
like to steal over us. From it, also, it is evident that the idea of the
Berleb. Bible, that these words are strictly suitable only in the lips of
Christ, is altogether erroneous. " Among us," it is there said, " no
man may, in his suffering, ask God why hast Thou sent this or that
affliction ? for we shall at all times find sufficient reason why we
have deserved this, and much more. All that a suffering man can say
is, 0 my, God forsake me notr The sufferer before us does not ask
why God, in general, allows him to suffer, but why He has forsaken
him. To this voliy, every one has a right, who can in truth call God
his God, notv/ithstanding his manifold failings. For " God has
forsaken no one who trusts in Him at all times," and God can forsake
no such one. In short, the expression, forsake me not, which alone,
it would appear, is admissible, is not essentially different from the
exclamation. Why hast Thou forsaken me, and must rest on
precisely the same ground. He only who can ask God, " Wherefore
hast Thou forsaken me?" can pray with confident assurance,
"Forsake me not." — The previous appellation, My God, my God,
contains the ground of the icherefore, the right to put such a
question. He who cannot call God his God, he who is without the
covenant and without the promises, he who has obtained no pledges
of the grace of God, may be justly forsaken : he has no ground to
implore of God, that He would show by the result that the desertion
is altogether a matter of appearance and feeling. Nay more, the
greater the right any one has to call God his God, the greater is the
confidence and decision with which he can utter tha why. Thus it is
evident that the most complete right to the why is reserved for one,
viz. Christ, who, in the full sense, can call God His God ; at the same
time, a sufficient right belongs also to all believers. The emphatic
repetition of the expression, Mv God, shows how firmly the sufferer
chugs to this his only groun^, of hope, how thoroughly conscious he
is that it is here that he is to find an antidote to despair, that it is
from this point that there must go forth a reaction against present
appearances. The expression, My God, occurring three times, here
and in ver. 2, VOL. I. 2 a
 370 THE BOOK OF PSALMS. is assuredly not accidental. —
The following remarks are Luther's : " Wherefore, let us shut up
these words in our hearts, and let us keep them carefully there, till
the proper time comes when we shall need them. Whoever cannot
comprehend them, let him remain with the people on the plain, in
the field below, and allow the disciples to go to Christ to the
mountain. Luke vi. 12, 17. For, not all the sayings of this Psalm are
uttered to each and every man, since all have not the same gifts,
and all have not the same sufferings. The Scriptures, according to
the circumstances of individuals, have milk for sucklings, and wine
and food for the strong ; so that there is consolation not only for the
weak, but also for the strong and for those who are endimufj 7 0 *D
great sufferings." — The second clause most interpreters, after the
example of the Septuagint and Luther (/ cry, hut my help is far\
translate : " far from my deliverance are the words of my
lamentation :" there is a great gulf between the cry for help, and the
help itself, which, now that matters are at the very last extremity
icith the sufferer — now that he stands with one foot in the grave, —
ought to stand in close contact with each other. Others translate : "
far from my help, from the words of my lamentation." This
translation is undoubtedly to be preferred. Were we to refer pim to
"•im, the plural would be required ; and, what is still more decisive,
the reference of pim to God is rendered necessaiy by the expression
pmn PX in the 11th and 19th verses. The cry in these verses, "be
not far," grows out of the address here, " Thou art fai'," after that
the impossibility of his continuing longer in existence had been
shown. God is far from the deliverance which He does not work out,
and from the complaint which He does not hear. This is all the more
painful, that the time for deliverance is just expiring, and that the
man from whom the complaint proceeds, is at the very gates of
death ; so that not to help now — not to hear now — appears to be
to give up altogether. We may not, however, adopt the view of most
of those who follow this exposition, and translate, "Thou art far."
This would require the pronoun : pim is in apposition to the pronoun
in ""jnaTy. The term nji^tj' signifies primarily, " roaring," or "
bellowing," and secondarily, " loud complaining." Ver. 2. My God, I
cry in the day time, and Thou answerest not ; and in the night time,
and I am not silent. Substantially, the "why" is to be supplied here
also. To be able to call God his God, and, in extreme distress, to cry
continually without
 rSALM XXir. VEE. 3. 371 being heard, is a striking
contradiction, wliicli imperiously calls for removal bj God's at length
hearing. The last words are translated by many, and I have no rest.
But the term T\'^ty\1 always signifies "silence;" and this translation
is particularly '; necessary here, in consequence of the opposition
between the i tei'm and the "-cry" of the first clause. The sufferer
can be ; silent when his cry finds an answer, when he gets assurance
of being heard and helped : so that thus / am not silent is exactly
parallel to Thou answerest not. Ver. 3. And Thou art holy, sitting
enthroned on IsraeV s 2^raise. There is no reason for substituting
and yet in room of the simple and ascertained and. The contrast
between the supposed reality and the idea — between the apparent
personal and the general experience — is not here indicated in
relation to the first and second verses, but is drawn for the first time
in vers. 6-8 in relation to the contents of vers. 3-5. The import is :
that I may lay down a further basis on which I rest my right to utter
the complaint, " Why hast Thou forsaken me? " — Tliou art holy, and
hast always taken an interest in Thine own people, hast never
forsaken any one of them ; but I appear to be altogether forsaken
by Thee. For Thou takest no interest in me, although I am note sunk
to the very depth of misery. — The idea of holiness in Scripture,
embraces in it the idea meant to be conveyed by theologians when
they define the term to be, " the highest purity in God demanding
the sanie purity on the part of the creature." This is evident from the
command, " Be ye holy, for I am holy ;" and Isa. vi. 5, where the
thrice repeated "holy" of the seraphims awakens in the prophet a
consciousness of his own impurity. But the two ideas are by no
means identical : the scriptural one is much more comprehensive
than the other. Holiness in the Scriptures comprehends majesty, as
well as holiness in the limited sense. God is holy, inasmuch as He is
separated from every created and finite being, and lifted above
them, particularly above sin, which can establish its seat only within
the domain of finite beings. The opinion of Gesenius (Thes.) and of
Kitzsch (Sys. 77), who would identify the scriptural with the
theological sense, is negatived by the very passage, the sixth
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