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VOLUME 1 NUMBER 1 FALL 1996
&XMPHIBLAN VSTION
CONSERVATION
THE INTERNATIONAL JOURNAL DEVOTED TO THE WORLDWIDE PRESERVATION
AND MANAGEMENT OF AMPHIBIAN AND REPTILIAN DIVERSITY —
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Amphibian Declines
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Rattlesnake Roundups
‘Herpetofauna in the Wildlif@lam
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What is BLUE CHAMELEON VENTURES?
Bill and Kathy Love operate the business based near
Ft. Myers in southwest Florida. We specialize
in providing quality photography for books, maga-
zines, calendars, web sites, signs, slideshows,
and all forms of commercial advertising. Our
area of expertise is herpetology - reptiles and am-
phibians, and also invertebrates. A huge stock of
color slides is available. Or, we'll be glad to dis-
cuss taking new shots of animals, displays, or
other subjects needed to illustrate your next
project.
Please call if we can help you with any photo needs.
Our slides have appeared in hundreds of publica-
tions over the past 17 years. Bill’s photos appear in
every issue of REPTILES magazine where he au-
thors a monthly question and answer column. He is
also a former partner of Glades Herp, Inc., a promi-
nent and respected breeder and supplier of
herpetofauna to zoos and collectors worldwide.
Other Aspects
We lead wildlife safaris to Madagascar, and also
captive breed many of the most beautiful spe-
cies of snakes and other herps. Please request
further information on both of these other as-
pects of our company.
NATURE
PHOTOGRAPHY
Tens of 1000’s of superb color
slides are available of reptiles,
amphibians and arachnids - all
aspects of their ecology, biol-
ogy, habitat, and captive main-
tenance and breeding. Query
us with your needs for use in
books, magazines, calenders,
as teaching/lecture aids, etc.
WORLD
ECO-TOURS
We offer guided adventure
tours/photo safaris to some
of the most unique and biologi-
cally interesting places on
Earth. Our specialty is the
unparalleled mini continent
of Madagascar. Contact us for
the brochures detailing our up-
coming “once-in-a-lifetime”
trips overseas.
HERPETOCULTURE
For over a decade, we’ve con-
centrated on selectively propa-
gating the most beautiful and
variable species in captivity.
Our emphasis has been on de-
fining and enhancing the color
and pattern traits that hold the
broadest appeal. Request our
annual breeder list.
A hungry Parson’s chameleon captures a
grasshopper with its long adhesive tongue.
wh x INE ssi BS
Matjaz Rojc and Michael Francis photograph-
ing a sleeping Leaftail Gecko on a recent tour.
Captive - bred hypomelanistic Honduran Milk
snake, a new inheritable trait that really glows!
ERRATUM SHEET (AMPHIBIAN & REPTILE CONSERVATION VOL 1 NO. 1)
To increase the accuracy of the journal and reduce the number of errors a new policy is now in place. This policy
was not followed in the first issue due to time constraints. All authors will now receive galley proofs once their
article has been edited and the layout completed. These proofs will be reviewed one last time by the author(s).
All errors should be brought to the attention of the editor as soon as possible.
The following journal corrections should be noted.
1) Page 3. Masthead.
a) The premiere issue (Vol. 1 No. 1) are now $12 each.
b) Back issues are $9 each (exception, Vol. 1 No. 1).
2) Page 10. In the Key Words section the word Viridi_ should be spelled Viridis.
3) Page 16. Under the photo caption the word Plethedon should be spelled Plethodon.
4) Page 20. The third sentence of the body of the article should read “Inbreeding depression may adversely
affect small populations by unmasking recessive deleterious alleles and reducing heterozygosity.”
5) Pages 24-26. Column.
a) The personal communication on page 25 (Groombridge 1994) was not attributed to Dr. Brian
Groombridge of the World Conservation Monitoring Centre (WCMC), Cambridge, England.
b) The WCMC database, referred to in the column, is in fact the CITES database of trade records.
WCMC have developed and maintain it on behalf of the CITES Secretariat.
c) By comparing export records and import records, and checking against known countries of origin, it is
in fact possible to get some idea of the extent of misreporting.
The up-dated information for this particular article was provided by Brian Groombridge on November 8, 1996.
NOTE - The most up-to-date Writer’s Guidelines and Manuscript Preparation instructions can always be found
at the Amphibian & Reptile Conservation website at: http://www.byu.edu/~arcon/
*
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AMPHIBIAN & REPTILE
CONSERVATION
THE INTERNATIONAL JOURNAL DEVOTED TO THE WORLDWIDE PRESERVATION
AND MANAGEMENT OF AMPHIBIAN AND REPTILIAN DIVERSITY
Volume 1, No. 1, Fall 1996
FOUNDER
Craig Hassapakis
Editor and p
Amphibian & Reptile Conservation
ADVISORY BOARD
Allison (
Center for Reproductio
oological Park
ian Institution
Departm:
Villanova L
Joseph T. Collins
Natural History Musewm
University of Kansas
Cari Gans
rtment of Biology
ty of Michigan
Dep
Uni
Howard E. Lawler
Executy ctor, Bio:
Tucson, Arizona
Roy W. McDiarmid
National B:
A Mittermeier
reation International
Washington, DC
George B. Rabb
Chairman, IUCN, Species Survival Commission
President, Chicago Zoological Society
Hobart M, Smith
Department of Environmental, Population
and Organismic Biology
University of Colorado
EDITORIAL REVIEW BOARD
C. Kenneth Dodd, Jr.
Southeastern Biological Scie
National Biological
e Center
rvice
Lee A. Fitzgerald
Department of Biology
University of New Mexico
Steven D. Garber
Wildlife Biologist
John F. Kennedy International Airport
Julian C. Lee
Department of Biology
University of Miami
Joseph C. Mitchell
Department of Biology
University of Richmond
Henry R. Mushinaky
Department of Biology
University of South Florida
Chri
Center for Envi
pher J. Razworthy
ental Research and Conservation
mbia University
Nelson Jorge da Silva. Jr.
Centro de Estudos ¢ Pesquisas Biologicas
Universidade Catolica de Goias, Brazil
Andrew T. Storfer
Center far Ecology, E' ion and Behavior
University of Kentucky
Robert J. Wiese
Assistant Director, Conservation and Science
American Zoo and Aquarium Association
CONTRIBUTORS
Section Editor
Amphibian Ecology & Conservation
Jamie K. Reaser
Center for Conservation Biology
Stanford University
Columnist
Allen Salzbery
New York Turtle & Tortoise Society
Neic York, Neto York
Zoo Liaison
Frank L. Slavens
Curator of Reptiles
Woodland Park Zoological Gardens
Editorial Assistants
Guenevere Nelson-Melby
David D, Adam.
Art Directors
Carin Young
Samuel D, Ashley
Amy Stout
Computer Support
David J. Owens
Printed on recycled paper with soy based ink
On the Cover
A male Golden toad (Bufo periglenes) at the
Monteverde Cloud Forest Reserve, Costa Rica.
Sadly, there have been no confirmed sightings of
Golden toads seen since May of 1989, when a
single male was seen. No one knows why this
species disappeared from its pristine forest sur-
roundings, far from human activities.
Golden toad photographs generously donated
by Michael and Patricia Fogden, Natural His-
tory Photographers, Kinbuck, Dunblane,
Perthshire FK15 9JU, Scotland, UK. Tel/fax: +44
1786 822069.
Contents
Features
4 The Elucidation of Amphibian Declines
Jamie K. Reaser
9 Rattlesnake Roundup in Kansas: A Brief History
David L. Reber and Allison Smith Reber
Scientific
15 Effects of Timber Harvesting Practices on Peaks of
Otter Salamander (Plethodon hubrichti) Populations
Joseph C. Mitchell, Jill. A. Wicknick, and Carl D. Anthony
Departments
20 Commentary
Population Biology and Herpetological Conservation:
A Cautionary Note
Andrew Storfer
24 Column
Herpetofauna in the Wildlife Trade and Nature: On the Difficulty
of Estimation
Allen Salzberg
27 News and Notes
29 Writer’s Guidelines and Manuscript Preparation
Editorial
28 Words from the Editor
Craig Hassapakis
Next Issue
Conservation of South African’s Endemic Dwarf
Chameleons
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© Amphibian & Reptile Conservation
Amphibian & Reptile Conservation (Fall 1996), 1(1): 4-9
Printed in the United States. All rights reserved
The Elucidation of Amphibian Declines
Are Amphibian Populations Disappearing?
Jamie K. Reaser
Center for Conservation Biology, Department of Biological Sciences, Stanford University, Stanford, Cali-
fornia 94305-1901
Key Words
Population declines, extinction, conservation, research, monitoring, amphibians
Abstract
Information regarding most am-
phibian declines is anecdotal and
_ natural fluctuations in amphib-
ian population size are not un-
common. However, biologists can
no longer find amphibians in re-
gions where they were once nu-
merous, and have directly ob-
served population declines and
species extinction. Inventory and
monitoring programs are being
established worldwide in order to
assess the status of amphibian
populations and to attempt to
identify causes of declines. Fac-
tors that may be contributing to
local amphibian declines include
natural population fluctuations,
natural succession and other
changes in vegetation, introduced
predators and competitors, patho-
gens, excessive collecting, toxic
compounds, and habitat destruc-
tion. Climate disturbance, pollut-
ants, and increases in UV-B ra-
diation have been implicated in
some well documented regional
amphibian losses. These factors
may decrease amphibian popula-
tion size by causing mass mortal-
ity, reducing the ability of indi-
viduals to produce viable off-
spring, and/or by inhibiting dis-
persal of individuals. A loss of
amphibians will have a significant
Correspondence, tel: (415) 725-9915,
fax: (415) 723-5920, e-mail: jkrtoad@
leland.Stanford.edu
impact on the state of the environ-
ment, as well as a decline in our
cultural heritage and human well-
being. Both biologists and con-
cerned citizens have vital roles in
amphibian conservation. A brief
list of possible citizen actions to
help protect amphibious life is pre-
sented here.
Amphibian populations world-
wide seem to be declining. Even
the causal observer can not find
frogs, toads, and other amphib-
ians as numerously as they once
could. Within the few short de-
cades of our lifetimes, a wide va-
riety of amphibians seem to have
been disappearing. Population
declines and species extinction
dot the pages of personal journals.
Biologists now search harder than
ever, asking “why” and hoping to
find the answers so that future
generations don’t have to be told
what they are missing.
Finding answers is no easier
than finding frogs. There is ample
evidence that humankind has
greatly impacted the distribution
and abundance of animal and
plant species worldwide through
extensive habitat alteration and
degradation. Such observations
can be made daily, and by non-
biologists. What we need to know,
however, is how specific land uses
impact the population dynamics of
amphibians, and at what spatial
and temporal scales. We also need
to know what the loss of amphib-
ians means for the continued func-
tioning of ecosystems. Ultimately,
we need to know what the disap-
pearance of amphibians signifies
for human well-being.
The paucity of data
Unfortunately, information re-
garding most amphibian declines
is anecdotal. For several species,
range reductions are well docu-
mented, but local population de-
clines are less evident. For most
species, studies only provide frag-
mentary pictures of populations
on population trends.
Research on amphibian ecol-
ogy has historically lagged behind
that of other vertebrate groups
because amphibians are often dif-
ficult to study and funding is
harder to obtain. Those concerned
about the loss of Neotropical mi-
gratory birds can reference broad
ranging, standardized datasets
from numerous monitoring pro-
grams. Several of these datasets
span multiple decades; one was ini-
tiated as early as 1900. However,
long-term (decade or longer) moni-
toring programs exist for only a
few amphibian species and only at
specific sites. Amphibian popula-
tion dynamics can typically be de-
scribed as “boom or bust;” natural
6 Jamie K. Reaser
fluctuations may be the rule rath-
er than the exception. Thus, only
very long-term datasets are use-
ful in validating suspected trends
and elucidating the mechanisms of
amphibian population declines.
The first worldwide effort to
assimilate data and hypothesize
the causes and consequences of
amphibian declines was held in
Irvine, California in 1990 (Blau-
stein and Wake 1990). Since this
meeting, an international investi-
gatory team, entitled the Declin-
ing Amphibian Populations Task
Force (DAPTF) of the Species
Survival Commission (SSC), In-
ternational Union for the Conser-
vation of Nature (IUCN), has been
formed and working groups have
been designated to address poten-
tial causes (e.g., toxins, UV-B ra-
diation, pathogens) and geogra-
phic regions. The findings of work-
ing groups and individual scien-
tists are published quarterly in
the DAPTF newsletter, Froglog,
making information readily avail-
able. Conservation organizations,
naturalist societies, and regional
agencies have been successful in
establishing local amphibian in-
ventory and monitoring programs
that often effectively utilize a mas-
sive volunteer work force. County-
and continent-wide initiatives,
such as the North American Am-
phibian Monitoring Program
(NAAMP), are in various stages of
design, testing, and implementa-
tion.
What we do know
Amphibian populations can fluctu-
ate greatly between years; varia-
tions in moisture, predation, com-
petition, disease, and catastrophic
events may greatly influence popu-
lation size. Populations may suffer
great losses. Yet, if the same popu-
lations experience “good years”
that result in many surviving off-
spring, the long-term population
trend may be stable. Long-term
stability may also be attained if
amphibians from other locations
recolonize sites where popula-
tions have been annihilated. For
example, Norman Weitzel and
Howard Panik observed Pacific
chorus frogs (Pseudacris regilla)
in Nevada and found that in 80%
of the years between 1975 and
1989 frogs produced offspring that
became members of the next gen-
eration. The population was anni-
hilated from the breeding pond ten
times by natural disturbances that
included flash floods, stream dry-
ups, and sudden increases in water
temperature. Yet, after each local
extinction event, this population
was soon reestablished by coloniz-
ing chorus frogs.
Extremely long-term data-
sets are required to distinguish be-
tween natural population fluctua-
tions and anthropogenically in-
duced declines. Joseph Pechmann
and his colleagues monitored
populations of one species of frog
and three species of aquatic-
breeding salamanders in the
southeastern U.S. for twelve con-
secutive years. They found no evi-
dence of drastic declines for any
species, although the populations
sizes did fluctuate. However, bi-
ologists Michael Reed and An-
drew Blaustein recently reana-
lyzed Pechmann’s data as well as
that from four other long-term
studies using a statistical tool
called power analysis. All these
studies, analyzed by this method,
indicated no declines. While the
datasets from these studies were
not extensive enough to reveal
statistical evidence of a decline,
the lack of decline in populations
of these amphibians could not be
supported. Thus, even with twelve
years worth of scientifically rigor-
ous data, the status of these am-
phibian populations cannot be de-
FEATURE (Scientific)
finitively assessed.
Amphibians cannot be found
in many of the locations where they
were once numerous. Yellow-leg-
ged frogs, red-legged frogs, spot-
ted frogs, leopard frogs, western
toads, cricket frogs, and tiger sala-
manders are a few of the North
American amphibians dwindling
in the number of sites of occur-
rence and population size. The am-
phibian queue for listing under the
United States’ federal Endan-
gered Species Act has become so
long that species ruled as justified
for protection are precluded from
it for years by stacks of preceding
paperwork.
Herpetologists have wit-
nessed the vanishing of amphibian
populations, and even entire spe-
cies. Biologists Stephen Corn and
James Fogleman conducted an ex-
ceptional study, documenting six
populations of the leopard frog
(Rana pipiens) in Colorado for the
decade 1973-1982. In 1973 only a
single population failed to repro-
duce. For frogs this may not be un-
usual, but by 1981 no leopard frogs
could be found at any of the sites.
The study ended with a total ab-
sence of R. pipiens in the region.
The golden toad (Bufo per-
iglenes) of Costa Rica, so named
because of the male’s bright or-
ange color, never failed to show up
for its annual spring breeding orgy
(see Figure 1, page 4) from the
early 1970s through 1987. Martha
Crump and her colleagues counted
1500 individual adult golden toads
in 1987, but noted that only 29 tad-
poles metamorphosed and joined
the population. From 1988 to 1990
these biologists located only 11
toads. Bufo periglenes has not been
seen at the study site since.
In Australia, just north of
Brisbane, a bizarre little frog was
discovered in 1973. The gastric
brooding frog (Rheobatrachus
silus), so named because it swal-
FEATURE (Scientific)
lowed and brooded its young in its
stomach, was an immediate won-
der to science and a potential boon
for physiologists interested in find-
ing cures for ulcers and possibly
other gastric disorders in humans.
A loss of worldwide significance,
the frog has not been seen since
1979, leaving little clue as to what
caused its extinction. Strangely
enough, its natural habitat was
found in seemingly pristine tropi-
cal forest, far from routine human
disturbance.
Why are amphibians
declining?
Could the above mentioned frogs
have succumbed to natural, local-
ized climatic disturbances such as
drought ... been the victims of op-
portunistic pathogens ... suffered
from a yet, undetected global at-
mospheric phenomenon? Could
the little known golden toad and
gastric brooding frog merely be
sitting it out underground, await-
ing what they consider more fa-
vorable conditions?
It is difficult to generalize as
to the causes of amphibian disap-
pearances. Not every amphibian
population, nor every species, is
declining. And, those that are de-
clining are doing so at varying
rates and scales. Figuring out
what is happening to amphibians
and why is exasperatingly difficult
and exhaustingly time consuming.
Locally, factors that may be
contributing to amphibian de-
clines include natural population
fluctuations, natural succession
and other changes in vegetation,
introduced predators and com-
petitors, pathogens, excessive col-
lecting, toxic compounds, and habi-
tat destruction. Climate distur-
bance, pollutants (particularly
those associated with acid deposi-
tion and pesticides), and increases
The Elucidation of Amphibian Declines 7
in UV-B radiation have been im-
plicated in some well documented
regional amphibian losses (see re-
views by Barinaga 1990; Wyman
1990; Blaustein and Wake 1990;
Tyler 1991; Phillips 1994; Blau-
stein and Wake 1995).
Unfortunately, there may be
a significant time lag between the
negative influence of a factor on
amphibians and evidence of a
population decline. Prospecting for
cause and effect relationships is,
therefore, exceptionally difficult
even in contemporary studies. Di-
agnosis is further complicated be-
cause factors can act in concert
and their relationship is rarely ob-
vious. For example, frogs have
been observed to die of an infec-
tion caused by a common, wide-
spread microbe called Aeromonas
hydrophilla that is not normally
pathogenic. Any number of other
factors may inhibit frogs’ immune
systems, making them susceptible
to infection.
Three general hypotheses
illustrate the mechanisms by
which various factors can cause
the extinction of amphibian popu-
lations, and eventually species:
1) Mass Mortality hypothesis
A factor or combination of factors
influences amphibians in such a
manner as to induce mortality of
individuals, sometimes entire
populations. Different factors
may contribute to mortality at
varying points in amphibian de-
velopment. However, the decline
of many populations is not merely
a problem of producing viable off-
spring. It is apparent that some
factors are influencing adult sur-
vivorship because many of the
rapid declines are occurring in
periods far shorter than the ani-
mals’ life span.
2) Reduction of Fitness
hypothesis
One or more sublethal factor(s)
reduces the ability of individual
amphibians to produce viable off-
spring (7.e., “fitness”). This eventu-
ally leads to population declines
and even population- and species-
level extinction. Genetic variation,
growth rate, size at maturity, lon-
gevity, and physiological con-
straints all influence the fitness of
amphibians. Some amphibians
have such specific conditions for
breeding that even subtle environ-
mental changes can result in the
failure of a population to breed.
3) Failure to Rescue
hypothesis
The observed declines are prima-
rily driven by the failure to rees-
tablish populations following lo-
cal extinction. Typically, when a
local population goes extinct, the
habitat is colonized by amphib-
ians dispersing from nearby
sources (this reestablishment is
termed the “rescue effect”). Under
this scenario, changes in the
chemical or structural environ-
ment prevent amphibians from
dispersing widely.
Several biological character-
istics of amphibians are likely to
impede recolonization following
local extinction: (1) physiological
limitations (particularly water re-
quirements) make it difficult,
even impossible, for amphibians
to persist in or travel through
suboptimal habitat; (2) amphib-
ians tend to have small home
ranges, many move only short dis-
tances, and rarely “wander”; and
(3) amphibians, especially the
adults of many species are ex-
tremely faithful to a specific loca-
tion, or set of locations, and are un-
likely to abandon sites even if they
can no longer breed there.
8 Jamie K. Reaser
The specific means (e.g.,
physiological processes) by which
these general mechanisms operate
are not well understood and are
rarely investigated. In part, this is
due to the fact that solving the
puzzle requires the cooperation of
experts across disciplines as di-
verse as geology and genetics. Re-
egretfully, most biologists are highly
specialized and rarely trained or
encouraged to work with col-
leagues from other fields. When
they have, however, pieces of the
puzzle fall into place. For example,
and interdisciplinary team in Or-
egon led by Andrew Blaustein is
now able to illustrate how strato-
spheric ozone depletion may lead
to amphibian population decline. It
works like this: Ultraviolet-b radia-
tion penetrates the Earth’s thin-
ning, protective ozone shield and
beams its way to earth where it
comes in contact with amphibian
eggs. The high-level and/or pro-
longed exposure to radiation caus-
es damage to eggs’ DNA (the ge-
netic information template) mol-
ecules, which in turn results in the
death of cells and thus tadpoles do
not develop. As adult frogs die and
are not replaced by new genera-
tions, the population declines and
eventually goes extinct. Blaustein
and the other investigators further
learned that different amphibian
species have varying amounts of
photolase, an enzyme that can re-
pair DNA damage. The declining
Cascades frog is low in photolase,
while the coexisting and successful
Pacific chorus frog has good DNA
repair capabilities; extra copies of
photolase genes secure protection.
The impact of amphibian
declines
A loss of amphibians will have a
significant impact on the state of
the environment. Amphibians are
vital components of the world’s
ecosystems. Amphibians comprise
one-quarter of all vertebrate spe-
cies on earth and sometimes con-
stitute the highest percentage of
vertebrate biomass in a given
area. This measure may be posi-
tively correlated with a species’
contribution to ecosystem func-
tion; 7.é., it is one indication of the
organisms’ importance to main-
taining the system’s integrity. Am-
phibians consume aquatic vegeta-
tion, invertebrates and other ver-
tebrates, and are eaten by numer-
ous predators. Therefore, amphib-
ians play multiple, vital roles in the
food chain of ecosystems.
Amphibians are apparently
declining even in seemingly pris-
tine, protected areas worldwide.
Because of these trends, many
biologists are pondering whether
amphibian declines should be in-
terpreted as a warning signal;
that is to say that the disappear-
ance of amphibians indicates that
something is gravely amiss in the
biosphere. Because amphibians
have permeable gills, skin, and
eggs; have diverse life histories;
are widely distributed and occupy
a variety of habitats, their popu-
lation dynamics may qualify as
reliable gauges of environmental
health Gf only we can learn to in-
terpret the signals).
Frogs are totems of luck for
numerous native culture; many
hunting poisons, ceremonial hallu-
cinogens, and medicinal drugs are
amphibian products. Amphibians
are chemical factories and the
compounds they produce may
hold cures to all sorts of ills, in-
cluding AIDS and cancer. If
you’ve had painkillers adminis-
tered recently, you may have a
frog or two to thank. For an excel-
lent review of amphibian contribu-
tions to medicine, see Grenard’s
(1994) Medical Herpetology.
FEATURE (Scientific)
You can make a difference
Approximately 5,000 amphibians
have been described by science,
with additional descriptions being
cataloged at a rate of 1 to 2 per-
cent a year. The rate of loss is
immeasurable; we don’t know
how many amphibians have come
and gone without recognition.
The amphibian decline “cri-
ses” demands that the status of
amphibian populations be rapidly
assessed and that where declines
are apparent, mechanisms be iden-
tified, managed, and recovery pro-
grams established. This is much
more easily stated than accom-
plished. There are far more am-
phibians than biologists investigat-
ing their declines. Funding is hard
to come by, particularly for the
long-term studies that are critical
to understanding amphibian popu-
lation dynamics. Also, time is not
on the side of the amphibian popu-
lation dynamics—human popula-
tion and resource consumption
continue to increase, rapidly
changing the landscape that am-
phibians have been evolving in for
roughly 350 million years.
Yes, there is hope. Amphib-
ian populations have rebounded
and sites have been recolonized
following massive die-offs. Main-
tenance and recovery of environ-
mental quality, and the restora-
tion of fragmented landscapes will
enable amphibians to persist.
As a citizen concerned about
amphibians, your role in amphib-
ian conservation is as critical as
that of any highly trained biologist.
The following is a very brief list of
the many actions that you can
take to help protect amphibians,
and maintain their vital roles in
the circle of life.
e Become a volunteer assistant for
a local amphibian monitoring pro-
FEATURE (Scientific)
gram or research project. Contact
your regional wildlife agency for
information on studies in your
area.
e Enlighten other people to the
wonders and plight of amphibians
by harnessing your enthusiasm
and knowledge. Talk to children,
the media, local officials, and the
voting public.
¢ Support legislation that pro-
motes healthy, intact ecosystems.
¢ Fight legislation that weakens
control of pollution and land de-
velopment.
¢ Encourage government agen-
cies to fund long-term research
projects on amphibians.
¢ Respect your wetlands by keep-
ing them healthy. Do not pollute
them with unnatural refuse such
as litter and harmful chemicals
(e.g., petroleum products and pes-
ticides).
* Organize routine cleanup pro-
jects.
¢ Admire amphibians in the wild;
don’t keep them as pets (animals
kept for research, in legitimate
conservation breeding projects,
and as educational displays such
as in zoological parks and aquari-
ums are not considered pets and
contribute to the conservation of
species).
By joining forces, biologists and con-
cerned citizens around the world
can become a very powerful lobby
for the conservation of amphibians.
And amphibians, inventoried and
monitored by these people, may be
a powerful gauge for ensuring the
protection of all life.
The Elucidation of Amphibian Declines 9
References
Barinaga, M. 1990. Where have all the
froggies gone? Science 24: 1033-1034.
Blaustein, A.R. 1994. UV repair and
resistance to solar UV-B in amphibian
eggs: A link to population declines?
Proceedings of the National Academy of
Sciences (USA) 91: 1791-1795.
Blaustein, A.R. and Wake, D.B. 1990.
Declining amphibian populations: A global
phenomenon? Trends in Ecology and
Evolution 5: 7.
Blaustein, A.R. and Wake, D.B. 1995. The
puzzle of declining amphibian populations.
Scientific American 272(4): 52-57.
Corn, PS. and Fogleman, J.C. 1984. Ex-
tinction of montane populations of the
northern leopard from (Rana pipiens)
in Colorado. Journal of Herpetology 18:
147-152.
Crump, M.L., Hensley, FR., and Clark,
K.L. 1992. Decline of the golden toad:
Underground or extinct? Copeia 1992:
413-420.
Grenard, S. 1994. Medical Herpetology.
Reptile and Amphibian Magazine, Potts-
ville, Pennsylvania. 139 pp.
Pechmann, J.H.K. and Wilbur, H.M. 1994.
Putting declining amphibian populations
in perspective: Natural fluctuation and
human impacts. Herpetologica 50: 65-
84.
Pechmann, J.H.K., Scott, D.E., Sem-
litsch, R.D., Caldwell, J.P, Vitt, L.J.,
and Gibbons, J.W. 1991. Declining am-
phibian populations: The problem of sep-
arating human impacts from natural
fluctuations. Science 253: 892-895.
Phillips, K. 1994. Tracking the Vanishing
Amphibians. St. Martin’s Press, New
York. 244 pp.
Reed, J.M. and Blaustein, A.R. 1995.
Assessment of “non-declining” amphibian
populations using power analysis. Con-
servation Biology. In Press.
Tyler, M.J. 1991. Declining amphibian
populations—a global phenomenon? An
Australian perspective. Altyes 9: 43-50.
Weitzel, N.H. and Panik, H.R. 1993.
Long-term fluctuations of an isolated
population of the Pacific chorus frog
(Pseudacris regilla) in northwestern
Nevada. Great Basin Naturalist 53:
379-384.
Wyman, R.L. 1990. What’s happening to
the amphibians? Conservation Biology
4: 350-352.
While the Bird Conservation Specialist
for the Smithsonian Institution’s Mi-
gratory Bird Center, Jamie K. Reaser
coauthored her first book, Bring Back
The Birds: What You Can Do To Save
Threatened Species (Stackpole Press
1995). In 1993 Dr. Paul Ehrlich, pre-
sented Jamie with the opportunity to
pursue her interest in amphibians
through formal investigation as his
doctoral student at Stanford Univer-
sity. Both Jamie and Dr. Paul Ehrlich
are interested in what amphibian pop-
ulation trends might indicate about the
health status of the biosphere.
One of Jamie’s greatest rewards of
her work with amphibians is the oppor-
tunity to mentor undergraduate stu-
dents who are interested in amphibian
ecology and conservation. An open in-
vitation is extended to those interested
to visit her at Stanford University.
© Amphibian & Reptile Conservation
Amphibian & Reptile Conservation (Fall 1996), 1(1): 10-14
Printed in the United States. All rights reserved
Rattlesnake Roundup in Kansas:
A Brief History
David L. Reber’ and Alison Smith Reber?
‘Assistant Director, Natural Heritage Center, Inc., 25124 Linwood Road, Lawrence, Kansas 66044-7311,
“Director, Natural Heritage Center, Inc. and Chair, Kansas Herpetological Society, Education and Conserva-
tion Initiatives, 303 West 39th Street, Hays, Kansas 67601-1518
Key Words
Rattlesnake roundups, commercialization, prairie rattlesnake, Kansas, conservation, legislation, poach-
ing, Crotalus viridi
Abstract
Organized hunting (“roundups”)
of rattlesnakes for commercial
profit has occurred for 50 or more
years in several states in the U.S.
For along period, such roundups
went unnoticed by wildlife agen-
cies, biologists, and herpetologists.
In recent years, there has been
increasing concern that such in-
tense and largely unregulated
hunting could have an adverse
effect on wild populations of
rattlesnakes. There have been few
conclusive studies on the topic.
The first organized rattlesnake
roundup to occur in Kansas was
held in September of 1992 in the
small town of Sharon Springs,
and it is the intent of this paper to
outline a series of events which
occurred in the first, formative
years of the roundup. Prior to the
introduction of rattlesnake round-
ups into Kansas, all native am-
phibians and reptiles were pro-
tected from commercial interests
by state law. The first roundup
had little economic benefit for the
participants, as all carcasses were
either given or thrown away. Be-
fore the second roundup (held less
than a year later) the organizers of
the roundup succeeded in changing
‘Correspondence, tel/fax: (913) 843-2205
(must contact in advance before send-
ing faxes), e-mail: reber@idir.net
state wildlife laws to accommodate
their economic interests. The ob-
jections and recommendations of
the Kansas Department of Wildlife
& Parks (KDWP), conservation
organizations, and professional
biologists were essentially ignored
so that a select few might profit
from commercial use of rattle-
snakes. It became clear that eco-
nomic gain for the roundup pro-
moters and vendors held a higher
priority than sound wildlife man-
agement. The legalized sale of
rattlesnakes and their parts, in-
cluding meat for human con-
sumption, raised new concerns
regarding meat inspection laws in
Kansas. Shortly before the third
roundup, the Kansas Department
of Health and Environment
(KDHE) informed the roundup
organizers that their practice of
selling uninspected meat was ille-
gal. Nonetheless, the sponsors sold
rattlesnake meat the following
weekend. Clearly, the roundup or-
ganizers intended to violate the law
as long as there was money to be
made. The roundup organizers
were never prosecuted and no fines
were levied. Although a rattle-
snake roundup has taken hold and
continues to prosper in Kansas, it
is clear that without the efforts of
biologists and concerned citizens
the situation would be much
worse. Many of the more mali-
cious aspects of the roundup have
been tempered as a direct result of
the efforts of relatively few indi-
viduals and organizations.
Word travels fast in the Kansas
Herpetological Society (KHS). All
were astounded and appalled that
the unthinkable was on the verge
of occurring right here in our very
own herpetologically enlightened
state. It can’t happen here. Bad
things, you know, always happen
somewhere else. Yet there, in gi-
ant black type on ugly electric yel-
low paper, was the announcement:
“Kansas’s First Ever Rattlesnake
Roundup!” It was to be held in
Sharon Springs, Kansas—spitting
distance from Colorado—on La-
bor Day Weekend 1992. We were
all too familiar with the broad-
ranging destructiveness of rattle-
snake roundups in other states:
the unregulated taking of wild ani-
mals for commercial profit, the use
of gasoline to flush snakes out of
shelters, the neglect and abuse of
captive animals, and the perpetu-
ation of antisnake sentiment. This
roundup was advertised as a
fundraiser; a family oriented event
with the added benefit of “control-
ling overpopulation” of rattle-
snakes which, sponsors claimed,
FEATURE (Popular)
cause immeasurable damage to
humans and domestic animals in
western Kansas.
Hosting a rattlesnake round-
up was not an idea that citizens of
Sharon Springs conceived on their
own. Members of the sponsoring
organizations had friendly ties
with James White, owner of the
“Fangs and Rattlers Show” from
Granbury, Texas. In fact, one wo-
man from Sharon Springs served
as a bridesmaid in White’s wed-
ding—a wedding that took place in
a snake pit. Sharon Springs
was looking for a fund-raiser,
and James White was looking
for another gig. The snakes,
they believed, were free for
the taking. A rattlesnake
roundup in Sharon Springs
was the perfect solution.
Herpetologists did not agree.
At the time, Kansas law
prohibited the sale of any wild
reptile or amphibian, includ-
ing prairie rattlesnakes, and
limited possession to five ani-
mals. The roundup sponsors
did plan a few tricks to bypass
these existing laws, such as
giving away rattlesnake
meat, provided that the re-
cipient purchased a bun on
which to eat it. Many herpe-
tologists concerned about the
rattlesnake roundups were confi-
dent that the state’s wildlife
agency would not stand for such
attempts to circumvent the law.
Several of these herpetologists
sent token letters of concern to
various individuals and agencies,
and returned responses ranged
from reassurance to hostility. Kan-
sas Herpetological Society (KHS)
representatives made several
vain attempts to educate roundup
sponsors and to shape the event
into something less destructive.
Nonetheless, the roundup com-
menced as planned.
An estimated 2000 people
Rattlesnake Roundup in Kansas 11
attended the roundup; among them
were KHS members Henry and
Virginia Fitch and Travis Taggart,
who had been contracted by the
Kansas Department of Wildlife &
Parks (KDWP) to collect data on
the size and reproductive condi-
tion of captured snakes. This data
collection alone provided little in-
formation about the effects of the
roundup. Other KHS members in
attendance to observe and report
on the activities of the roundup
included Dr. David Edds of Empo-
on
One of the sideshow type tricks shown at the
“Fangs and Rattler” show.
ria State University, Randall Reis-
erer, and David Reber of the Uni-
versity of Kansas. Several veter-
ans of Oklahoma’s roundups were
also in attendance, to help in the
“snake pit” and to guide groups in
search of prairie rattlesnakes. A
total of 75 prairie rattlesnakes
were turned in (Edds 1992), 18 of
which were born at the roundup.
All 75 were ultimately killed. The
snake meat sales trick failed, and
most of the meat was either given
away or thrown away, as were all
other snake parts. The roundup did
earn a substantial amount of mon-
ey, through assorted trinket ven-
dors, snake products from other
states, food service, and tickets to
the “Fangs and Rattlers Show,”
direct from Granbury, Texas. The
“Fangs and Rattlers Show” was
typical roundup entertainment,
the announcer cajoling the crowd
while the pit crew performed dare-
devil tricks, most of which involved
unnecessary rough treatment of
the animals. Children were invited
to pet loosely restrained rattle-
snakes, and dangle inflated bal-
loons into the pit in hopes of entic-
ing a strike. Overall, a carnival-
like atmosphere pervaded
and, with current laws prohib-
iting the sale of wild caught
prairie rattlesnakes, the
snakes themselves were not
the central money maker.
Those of us in attendance at
the roundup remained confi-
dent that the roundup organiz-
ers would lose interest and
move on to something less de-
structive. The thought that ex-
isting laws could be legislated
moot never crossed our minds.
Enter Sheila Frahm,
Senate majority leader repre-
senting the Sharon Springs
area. Senator Frahm began
her political career as a mem-
ber of the Sharon Springs lo-
cal school board. Unknown to
us, Senator Frahm had also at-
tended the first roundup, even
helping to weigh the snakes. Then,
during the 1993 legislative ses-
sion, she introduced a bill (Senate
Bill 137) that, if passed, would not
only allow the sale of prairie rattle-
snakes, but would raise the limit
from 5 to 30, and remove all man-
agement authority from the De-
partment of Wildlife & Parks.
Once again we thought: it can’t
happen here. We all assumed that
our enlightened legislature would
never pass such an anticonserva-
tion bill. Still, KHS and numerous
herpetologists provided testi-
mony against the bill, as did the
DAVID L. REBER
12 David L. Reber and Alison Smith Reber
Kansas Wildlife Federation, the
Kansas Audubon Council, the To-
peka Zoo, the Sedgwick County
Zoological Society, the Kansas
Chapter of the Sierra Club, and
even the Kansas Department of
Wildlife & Parks (KDWP). Appar-
ently politics, not prudence, drove
the issue, as the bill passed with
little opposition. There was some
success for conservationists; the
final bill was altered to place
regulation of the roundup back
into the hands of KDWP.
Before KDWP could enact
regulations, a second
roundup was held in the
spring of 1993, less than
one month after Senate
Bill 137 became law. A
springtime roundup was
recommended by repre-
sentatives of Oklahoma
and Texas roundups. This,
combined with the gener-
ous bag limit allowed by
the legislature, resulted in
over 170 snakes being
taken—more than double
the number of the first
roundup (Western Times
1993). Senator Frahm and
her daughter Chrissy were
in attendance and, in recognition
of her “heroic” efforts, Frahm was
presented with her very own
snake bucket and tongs.
Shortly after the second
roundup, KDWP began the long
process of developing regulations
for commercial use of prairie
rattlesnakes. Initially, KDWP
asked Oklahoma officials for ad-
vice on the issue, in spite of the
fact that Oklahoma roundups tar-
get a different (and much more
prolific) species, the western dia-
mondback. Roundup organizers
lobbied KDWP, asking for no pos-
session limit, a year-round season,
and no restrictions on the open
area. KHS (1993) president Dr.
David Edds attended virtually ev-
ery meeting of the Wildlife &
Parks Commission, recommend-
ing that KDWP reset the posses-
sion limit at 5 snakes, limit the
open area to only the counties
surrounding Sharon Springs, and
limit the open season to the round-
up weekend. Many others ad-
dressed the Commission as well,
advocating moderation in light of
the glaring lack of sound biologi-
cal information. In spite of their
testimony, each draft of the regu-
lations proved more lenient. Fur-
thermore, when herpetologists at-
Another one of the sideshow type tricks shown at the
“Fangs and Rattler” show.
tended, the issue was often re-
moved from the meeting agenda,
as if the Commission were at-
tempting to push regulations
through without input from the sci-
entific community. Again, politics
drove the issue.
It was decided by the KHS
that their position statement
must be reinforced using the avail-
able scientific literature and pub-
lished, for permanent record and
to assist policymakers. Thus, a po-
sition paper was written, totaling
11 pages and 48 references. The
paper included recommendations
for regulating the roundup—rec-
ommendations aimed at pro-
moting sportsmanship and a sus-
tainable yield of Crotalus viridis.
FEATURE (Popular)
A copy of the paper was provided
to each member of the Wildlife &
Parks Commission, as well as to
the Department Secretary, Theo-
dore Ensley. At the final ruling in
January of 1994, conservationists
were extremely disappointed. The
lenient draft regulations passed
with few modifications. The only
successes were lowering the pos-
session limit from 30 to 20, limit-
ing the open season to a 30 day
period prior to the roundup, and
limiting the open area to the west-
ern third of the state (approxi-
mately one half of the prai-
rie rattlesnake’s range in
Kansas). One major loop-
hole remained: the spon-
soring organizations were
not subject to a possession
limit. Thus, one person
could conceivably catch
their limit, sell their catch
to the roundup sponsors,
then head back for more,
effectively circumventing
the existing possession
limit. KDWP defended the
regulations as a compro-
mise between interest
groups.
Clearly, arguments based on
the biology of the prairie rattle-
snake and sound wildlife manage-
ment were not taken seriously. The
bottom line was money, and a way
to touch the bankbooks of the
roundup sponsors was badly needed
if their plans were to be altered.
KHS discovered that the Kansas
Department of Health and Environ-
ment (KDHE) has strict regulations
pertaining to the butchering and sale
of meat. Essentially, both the butch-
ering facility and the meat itself
must be inspected and passed by
KDHE before any meat can be sold.
A letter was sent by KHS to KDHE
immediately, along with an expla-
nation of the past butchering pro-
cess and an advertising flyer for the
upcoming roundup. KDHE repre-
DAVIDL REBER
FEATURE (Popular)
Rattlesnake Roundup in Kansas 13
a
possibly fashioned into an item such as a hatband).
sentatives contacted the roundup
organizers, explained meat pro-
cessing and sales laws, and in-
formed them of the penalties for vio-
lating those laws.
The third roundup was held
in May of 1994. The take totaled
over 300 animals, again doubling
the previous year’s take (Fitch
1994). KDWP’s research indicated
most of these snakes had been
stockpiled for extended periods,
despite the open season having
been only one month. Virtually ev-
ery snake exhibited dermal rot-
ting, lesions, abrasions, lacera-
tions, or paralysis, and most were
severely emaciated (Taggart
1994). Some were dead, presum-
ably having been crushed under
the accumulated mass of snakes in
small containers. Owing to the
poor condition and small size of
the prairie rattlesnakes, and the
tediousness of butchering small,
tough-skinned snakes, the round-
up sponsors purchased a number
of western diamondback rattle-
snakes to butcher and sell. The
butchering facility, located in a
livestock wash area at the local
fairgrounds, had been cleaned up
a bit (members of the audience
were no longer invited to ceremo-
niously behead snakes with a
rusted hatchet, for example);
however, they continued to ille-
gally butcher and sell uninspected
meat. The violations were re-
ported to KDHE. In spite of prior
warnings, the roundup sponsors
pleaded ignorance and received
only a letter of reprimand—no fine
was levied. KDHE officials did
state, however, that a second of-
fense would be prosecuted.
The Kansas Herpetological
Society has always encouraged
conservation of amphibians and
reptiles in Kansas. Laws protect-
ing wildlife from commercializa-
tion had provided a strong foun-
dation for these efforts, but the
advent of the rattlesnake roundup
Prairie rattlesnake (Crotalus viridus) which was taken at a roundup in 1992 (it can be assumed that it was slaughtered—
has weakened that foundation con-
siderably. It will be an arduous
task to regain what we once had,
but it can be done. Efforts to date
have resulted in many significant
improvements, and have buffered
many of the more negative as-
pects of the roundup. Some of the
more malicious aspects of the first
roundup have been diminished or
eliminated altogether. For ex-
ample, at the first roundup the
butcher shop offered spectators a
chance to swing the hatchet. This
was done ceremoniously, and was
portrayed as insidious revenge on
the rattlesnake. By the third
roundup, the “ceremony” was
gone, and the butchering was
nothing more and nothing less
than butchering. Direct negative
effects on children have also been
reduced slightly. At the first
roundup, children were invited to
participate (for a fee, of course) in
the torment of rattlesnakes via
“balloon fishing”. Children waved
DAVID L REBER
14 David L. Reber and Alison Smith Reber
inflated balloons over the heads of
rattlesnakes while the pit atten-
dant stomped at the snakes in an
effort to elicit a strike. The “bal-
loon fishing” activity has now been
eliminated. Furthermore, the
roundup sponsors no longer ad-
vertise the event as a means of
“controlling overpopulation” of
rattlesnakes. Nor do they continu-
ally deride the animals as the
scourge of the prairie. Overall,
these small improvements add up
to an event that is much less per-
nicious to young minds than it
might have been. However, there
have been some less-than-honest
attempts to pacify the conserva-
tion community. For the third
roundup, the sponsors had desig-
nated several areas totaling 88
square miles as “rattlesnake pre-
serves” where hunting of rattle-
snakes was not allowed. Further
investigation revealed that these
areas were nothing more than
land for which the owners had de-
nied permission to trespass. Thus,
the roundup sponsors did not in
fact designate “rattlesnake pre-
serves,” but simply capitalized on
inaccessible land as a public rela-
tions gimmick.
Some headway has been
made with respect to regulating
the roundup. Although a 20 snake
limit is still far too high, and a 30
day season far too long, it is likely
that, without the efforts of many
herpetologists and conservation-
ists, the sponsors would have been
granted their request of no posses-
sion limit and a 12 month season.
In addition, the entire state of
Kansas would likely have been
opened to the roundup, despite the
fact that prairie rattlesnakes only
live in the western half. Having no
limit on where rattlesnakes could
be collected throughout the state
could have placed other species,
including the state-protected tim-
ber rattlesnake, at risk. Much has
been accomplished, yet there is
still much to be done.
Several changes have oc-
curred since the third roundup.
Senator Frahm is now Lieutenant
Governor Frahm. Because she has
lost her powerful position in the
legislature, she has much less abil-
ity to persuade other legislators,
and less power also with the
KDWP. In addition, she is now
subject to more intense scrutiny
than in the past, and thus she may
be less apt to promote anticonser-
vation measures. Also, there is
some question about the location of
the fourth roundup, as it is ru-
mored it will move to the nearby
town of Colby, Kansas, the home-
town of Sheila Frahm. Whether
these changes are good or bad is
largely up to the conservation
community, and how it acts con-
cerning them. There are several
trump cards yet to be played, but
in the long run it is the grassroots
effort that will keep the snakes in
the grass—where they belong.
References
Edds, D. 1992. Observations on the 1992
Sharon Springs Rattlesnake Roundup.
Kansas Herpetological Society News-
letter, No. 90.
Fitch, H.S. 1994. The Sharon Springs
rattlesnake roundup. April 29-May 1,
1994. Unpublished report to Kansas
Department of Wildlife & Parks. 10
pages.
Taggart, T. 1994. Letter to Robert F.
Hartmann, Investigation and Inventory
Supervisor, Kansas Department of
Wildlife & Parks. Kansas Herpetol-
ogical Society Newsletter, No. 98.
Western Times, The. 1993. Large
Crowd Attend Rattlesnake Hunt. The
Western Times, May 6, 1993.
FEATURE (Popular)
David L. Reber earned a BS in Systemat-
ics and Ecology from the University of
Kansas in 1991. He then pursued grad-
uate work at the University of Kansas
in the School of Education, and in 1995
received a Kansas Teaching Certificate
for middle and secondary level science.
During his graduate work, David was
voted president-elect of the Kansas Herpe-
tological Society (KHS), of which he
had been a member since 1980. During
his three years on the KHS executive
council, he has focused the society’s
efforts on conservation issues includ-
ing, but not limited to, rattlesnake
roundups. He now teaches at the Natwr-
al Heritage Center, Inc., where he is also
associate director, and at Raintree Ele-
mentary School in Lawrence, Kansas.
In 1985, he saw a photograph of a
young field biologist with four bull-
snakes in her lap; he later married her.
Alison L. Reber earned a bachelor’s
degree in Environmental Studies from
the University of Kansas in 1994. She
has been an active member of KHS, is
coauthor of the KHS Position Paper
Regarding Rattlesnake Roundups, and
has coordinated many efforts to reduce
the impact of commercial taking of prat-
rie rattlesnakes in Kansas. Alison has
provided hundreds of children with
hands-on natural history experiences,
often emphasizing herpetology. In early
1995, she founded the Natural Heritage
Center, Inc. (NHC), a nonprofit chil-
dren’s science education center in
Lawrence, Kansas, which she now di-
rects. She also teaches at Raintree
Montessori School in Lawrence. Alison
is a firm believer in the power of edu-
cation. Both authors hope that early,
positive experiences with nature will
help people make wiser decisions re-
garding our natural resources.
For further reading about rattle-
snake roundups the authors suggest the
Kansas Herpetological Society Position
Paper Regarding Rattlesnake Round-
ups, (Kansas Herpetological Society
Newsletter, No. 96: 9-20, 1994), The
Sweetwater Rattlesnake Roundup: A
Case Study in Environmental Ethics, by
Jack Weir (Conservation Biology, Vol-
ume 6, Number 20), and Driving Out
the Dread Serpent, by Ted Williams
(Audubon, September 1990).
ry
© Amphibian & Reptile Conservation
i Amphibian & Reptile Conservation (Fall 1996), 1(1): 15-19
Printed in the United States. All rights reserved
Effects of Timber Harvesting Practices on
Peaks Of Otter Salamander (Plethodon
hubrichti) Populations
Joseph C. Mitchell’, Jill A. Wicknick?, and Carl D. Anthony’
‘Department of Biology, University of Richmond, Richmond, Virginia 23173-0002, °>Department of Biol-
ogy, University of Southwestern Louisiana, Lafayette, Louisiana 70504, *Department of Biology, Univer-
sity of Southwestern Louisiana, Lafayette, Louisiana 70504
Abstract
The Peaks of Otter salamander (Plethodon hubrichti) 7s endemic to a small portion of the Blue Ridge
Mountains in Virginia. Much of its range lies within a high timber producing area owned by the National
Forest Service. Comparisons of salamander abundance on replicated transects in recent clearcuts, older
clearcuts, recent shelterwood sites, and mature sites revealed no significant differences. However, recent
clearcuts, supported consistently fewer salamanders than other sites. Salamanders in mature sites con-
sumed significantly more soft-bodied prey than in other sites. Numbers of hard-bodied prey did not differ
among sites. Timber harvesting practices do not eliminate this species but may diminish population size
and diet quality.
Key words
Plethodon hubrichti, timber harvesting, populations, Peaks of Otter salamander, conservation, ecology,
natural history, Virginia
Introduction
The entire range of the Peaks of Otter salamander
(Plethodon hubrichti) is limited to an approximately
19 kilometers (km) long portion of the Blue Ridge
Mountains in Bedford and Botetourt Counties, Vir-
ginia (Highton 1986; Pague and Mitchell 1991). Its
habitat is limited to elevations above 443 meters (m)
in deciduous forest and densities are highest in ar-
eas containing mature hardwoods. Logging impact
on terrestrial salamanders has resulted in complete
extirpation of local populations of other terrestrial
plethodontids and in population fragmentation with
probable genetic and demographic consequences
(Ash 1988; Buhlmann, et al. 1988; Dodd 1989, 1991;
Petranka, et al. 1993, 1994). If long-term conserva-
tion of the Peaks of Otter salamander is to prevail
it will require that populations not be severely im-
pacted by timbering practices that lead to local ex-
tinctions.
Several kinds of timber operations (clearcuts,
Correspondence, tel: (804) 740-7086, fax: (S04) 289-8233,
e-mail: mitchell@urvax.urich.edu
shelterwood cuts, group selection cuts) and poten-
tial, defoliation by gypsy moths may affect the
salamander’s forest floor habitat. Drying of leaf lit-
ter and humus layers due to canopy removal
(Pough, et al. 1987; Dodd 1991; Dupuis, et al. 1995)
limits salamander movements and the ability to for-
age. Jaeger (1990) and Jaeger and Barnard (1981)
clearly show that Red-backed salamanders (Pleth-
odon cinereus) foraged less and consumed fewer prey
in dry periods than in wet periods. If the canopy ina
logged site is eliminated and the forest floor becomes
relatively drier (as compared to a forested site), then
we would expect salamanders to be able to forage
less often and obtain fewer prey. Lowered prey con-
sumption may affect other aspects of their life his-
tory. Assessments of the impact of timber harvest-
ing practices on terrestrial salamanders such as the
Peaks of Otter salamander are necessary for biolo-
gists and resource managers interested in its con-
servation and in the economic uses of the forest.
In this preliminary report on a multiyear study,
we address the following objectives: (1) To compare
the size of P. hubrichti populations in sites that have
received four different types of timber management,
16 Joseph C. Mitchell, et al.
SCIENTIFIC
Table 1. Average numbers (+/- one standard deviation and range) of Peaks of Otter salamanders in replicates of three types
of timber management and mature forest stands in fall 1994 and spring 1995.
Treatment No. Replicates Fall 1994
Recent clearcut 6 4.7 +/- 4.6 (0-12)
Older clearcut 6 8.0 +/- 9.2 (0-25)
Shelterwood 5 8.0 +/- 10.8 (1-27)
Mature 6 8.8 +/- 5.0 (1-15)
Overall 7.4 +/- 7.7
and (2) To elucidate prey use patterns of populations
in the various treatments.
Materials and Methods
A total of 23 transects were established 16-18 June
1994 in the following treatment categories: recent
(4-5 years) clear cuts (number (n) = 6), 12-18 year
old clearcuts (n = 6), 2-4 year old shelterwood cuts
(n = 5), and mature (>80 years old) hardwood sites
(n = 6). There were too few separate shelterwood
cuts in the area to obtain a sixth site.
At each site, we established a 1 x 50 m tran-
sect by flagging woody vegetation and by placing wire
flagging in the ground at every 10 meters. Each tran-
sect remained permanent from season to season,
= coal FAS **
A topotype specimen of a Peaks of Otter salamander (Plethedon hubridhti). Bedford County, Virginia.
Spring 1995
4.0 +/- 3.8 (0-8)
3.7 +/- 4.0 (0-9)
3.2 +/- 2.9 (0-7)
7.3 +/- 5.8 (3-17)
4.6 +/- 4.3
except for two that were vandalized during the win-
ter of 1994-1995. These were reestablished in the
exact locations of the original transects in spring
1995. All transects were separated from each other
by a different type of stand, roads, or a distance >100
meters.
Searches for Peaks of Otter salamanders were
conducted in fall 1994 (12 September to 17 October)
and spring 1995 (9 May to 3 June) at night during or
just after rain when the forest floor was wet.
Transects were selected in random order each night
surveys that were conducted and each was walked
slowly by 1-2 people using headlamps. All sala-
manders during the spring sampling season were
released at their capture locations within a few min-
utes; in the fall sampling period they were released
o , -
WAYNE VAN DEVENDER
SCIENTIFIC
*. 5 . 5 =, y =e x 5 SH
= =. ye yo ze q : oes
= TSS ese Se = ss
A Peaks of Otter S
several days later (see below).
During the fall sampling period, we obtained
stomach contents by stomach flushing salamanders
(Fraser 1976). Prey were preserved in alcohol in in-
dividually labeled vials for later analysis. All sala-
manders were released within 5 m of their original
capture locations 2-4 days after collection. In the
laboratory, we identified 96% of prey items to order
and where possible, to family following Borrer, et
al. (1989). Remaining prey were identified to phylum
or class.
Results
Population sizes varied among stand types and be-
tween the fall and spring seasons (Table 1). Numbers
of salamanders ranged from 0 at a recent clearcut site
to 27 at a shelterwood site. In the fall, the average
number of salamanders in recent clearcuts was nearly
half that in the other stand types. However, the wide
variation in numbers of salamanders within stand
types, especially for shelterwood cuts, resulted in no
significant differences (ANOVA, F = 0.35, P = 0.792).
The spring assessment yielded fewer sala-
manders compared to the previous fall season (Table
1). The spring trend in numbers per stand type was
similar to that for the fall sample except for the lower
counts in the shelterwood and older clearcut sites.
The average number of salamanders within mature
stands was higher than those in clearcuts and
shelterwood cuts (Table 1). However, the variation
in numbers of salamanders in stand types did not
differ significantly (ANOVA, F = 1.10, P = 0.372).
We found 949 taxonomically identifiable prey
items in the stomachs of 80 salamanders from 20
sites. Ants (Hymenoptera) and collembolans
(Collembola) made up 54.6% of all individual prey
items in salamander stomachs. Of these, ants com-
prised 32.2% of the sample and collembolans 67.8%.
Effects of Timber Harvesting Practices on Salamander Populations
alamander (Plethodon hubrichti). Bedford County, Virginia.
IT
WAYNE VAN DEVEND!
We compared the numbers of ants and collembolans
separately among the four stand types with
Kruskal-Wallis tests. There was no significant dif-
ference in the numbers of ants (hard-bodied prey,
Jaeger 1990) consumed among the four stand types
(H = 2.102, df = 3, P = 0.552) (Figure 1a). In contrast,
there was a significant difference among the num-
bers of collembolans (soft-bodied prey, Jaeger 1990)
consumed (H = 16.794, df =3, P < 0.001) (Figure 1b).
Multiple comparison tests showed that salamanders
from mature stands ate more collembolans than sala-
manders from the old clearcuts (z = 2.68, P < 0.05)
and the shelterwood cuts (z = 3.98, P < 0.05).
Mean no. of ants per stomach
recent shelterwood
clearcut clearcut cut
sy ey
Nn ff OO
Mean no. of collembolans per stomach
o
recent old mature shelterwood
clearcut clearcut cut
Figure 1. Average number of ants (a) and collembolans (b) per
stomach in Peaks of Otter salamanders in four forest stand types.
The distribution of the letters refers to statistical results (see text).
18 Joseph C. Mitchell, et al.
Discussion
Population sizes of terrestrial salamanders are vari-
able and depend on a wide range of factors. These
include soil depth, soil temperature, soil moisture,
aspect, slope angle, underground shelter availabil-
ity, nest site availability, number of surface cover
sites for territories, prey quality and abundance,
predator abundance, and presence of known competi-
tors (Plethodon cinereus) (Buhlmann, et al. 1988;
Dodd 1991; Wicknick 1995). Clearing of the canopy
vegetation and the majority of understory trees, a
common result from clearcutting, changes the physi-
eal environmental characteristics (e.g., soil and log
moisture) of the area (Heatwole 1962; Blymyer and
McGinnes 1977; Dodd 1989, 1991; Dupuis, et al. 1995).
Loss of individual salamanders occurs directly from
logging operations that include road building, the use
of skidders and other heavy equipment, and mechani-
cal site preparation (Dodd 1991). Individuals not di-
rectly impacted by the immediate logging operation
are probably subjected to stresses associated with
reduced or altered prey resources and changes in the
physical characteristics of the soil/litter ecosystem.
We would thus expect to find reduced numbers of
Peaks of Otter salamanders in such areas.
Shelterwood cuts allow a possible solution to
the problems attached to clearcutting. Unfortu-
nately, shelterwood operations that leave a small
basal area (7.e., as few canopy trees) act identically
to clearcuts and they produce similar effects on sala-
manders. Our data show that the numbers of Peaks
of Otter salamanders in shelterwood cuts ranged
from one to 27, the largest range of variation in any
of the stand types. The numbers of Peaks of Otter
salamanders in shelterwood sites were, on average,
10-66% lower than in adjacent mature sites. The
wide variation may be related to the number of
standing trees remaining. Thus, different levels of
shelterwood cuts may have dramatically different
effects on Peaks of Otter salamanders because of the
interaction between the amount of basal area re-
maining and the quality of the habitat present be-
fore and after the operation takes place.
Population sizes in our study were, on aver-
age, consistently higher in mature sites that had not
been logged in 80 or more years when compared to
recent and older clearcuts and shelterwood cuts.
The high variation in number of salamanders within
stand types complicates the interpretation of these
data. The lack of statistical significance does not
SCIENTIFIC
mean that there are no detrimental effects caused
by these logging practices. Such effects may not be
detectable at the population level because of histori-
cal factors (e.g., past logging history and habitat qual-
ity), and differences in relative abundances due to
slope angle and aspect. The size of the impacted area
and its proximity to mature stands containing large
populations may influence the length of time for P.
hubrichti populations to achieve prelogging levels.
The effects of logging may be more clearly elu-
cidated by analysis of diet quality. The quantity of
collembolans consumed by Peaks of Otter sala-
manders was significantly higher in mature stands
than in recent clearcuts and shelterwood cuts. Col-
lembolans are soft-bodied prey (Jaeger 1990) which
presumably pass through P. hubrichti digestive
tracts quickly and with high assimilation efficiency,
as do other soft-bodied prey such as termites in the
congener P. cinereus (Gabor and Jaeger 1995). Dif-
ferences in diet quality among stand types suggests
that there may also be differences in salamander
growth and reproduction. Mature sites, therefore,
appear to offer a higher quality habitat to Peaks of
Otter salamanders than timbered sites because they
presumably have a more intact and functional soil/
litter ecosystem due to the types and quality of
downed woody debris, canopy shelter that affects
the thermal and moisture regime, and higher prey
quality. Results from our forthcoming analyses of
data on prey quality and availability in wet and dry
years will provide a detailed assessment of differ-
ential affects of stand types at the individual level.
Conclusions
The preliminary results of our study allow several
tentative conclusions. Peaks of Otter salamander
populations are not always completely eliminated
from a site within their range by timber operations
of clearcutting and shelterwood cutting. They are
reduced 45-47% by clearcutting and 10-66% by
shelterwood cutting, as compared to populations in
adjacent mature sites. Salamanders in mature sites
may obtain a higher quality diet than those in sites
treated by some form of timber management. Peaks
of Otter salamander populations remain at varying
levels of risk from timber management depending
on the type of harvesting practice used. Such prac-
tices undoubtedly cause small scale geographic
variation in growth, diet, reproduction, and popu-
lation recruitment.
SCIENTIFIC
Acknowledgments
We thank John Bellemore, Fred Huber, Larry Neuhs, and
Glen Szarzinski for facilitating our research in the George
Washington and Jefferson National Forest. Paul Sattler and
Gordon Wilson assisted in the field. For the analysis of
stomach contents, we thank Darryl Felder for laboratory
space and equipment and Caitlin Gabor and Sergio Nates for
references. Ken Dodd provided a thorough review of the
manuscript. This study is funded by grants from the George
Washington and Jefferson National Forest and the U.S. Fish
and Wildlife Service. Jill Wicknick and Carl Anthony were
also supported, in part, by NSF grant #DEB-9314081 to
Robert G. Jaeger.
References
Ash, A.N. 1988. Disappearance of salamanders from clearcut
plots. Journal of the Elisha Mitchell Scientific Society 104:
116-122.
Blymyer, M.J. and McGinnes, B.S. 1977. Observations on
possible detrimental effects of clearcutting on terrestrial
amphibians. Bulletin of the Maryland Herpetological Society
13: 79-83.
Borrer, D.J., Triplehorn, C.A., and Johnson, N.F. 1989. An
Introduction to the Study of Insects, 6th edition. Saunders
College Publishing, Philadelphia, Pennsylvania. 875 pp.
Buhlmann, K.A., Pague, C.A., Mitchell, J.C., and Glasgow,
R.B. 1988. Forestry operations and terrestrial salamanders:
Techniques in a study of the Cow Knob salamander, Plethodon
punctatus, pp. 38-44 in Szaro, R.C. et al. (editors). Manage-
ment of Amphibians, Reptiles, and Small Mammals in
North America. USDA Forest Service General Technical
Report RM-166. 458 pp.
Dodd, C.K., Jr. 1989. Status of the Red Hills salamander is
reassessed. Endangered Species Technical Bulletin 24: 10-11.
Dodd, C.K., Jr. 1991. The status of the Red Hills salamander,
Phaeognathus hubrichti, Alabama, USA, 1976-1988. Biological
Conservation 55: 57-75.
Dupuis, L.A., Smith, J.N.M., and Bunnell, F. 1995. Relation
of terrestrial-breeding amphibian abundance to tree-stand
age. Conservation Biology 9: 645-653.
Fraser, D.F. 1976. Empirical evaluation of the hypothesis of food
competition in salamanders of the genus Plethodon. Ecology
57: 458-471.
Gabor, C.R. and Jaeger, R.G. 1995. Resource quality affects
the agonistic behaviour of territorial salamanders. Animal
Behaviour 49: 71-79.
Heatwole, H. 1962. Environmental factors influencing local
distribution and activity of the salamander, Plethodon cinereus.
Ecology 43: 460-472.
Effects of Timber Harvesting Practices on Salamander Populations 19
Highton, R. 1986. Plethodon hubrichti. Catalogue of Amer-
ican Amphibians and Reptiles. 393.1-393.2.
Jaeger, R.G. 1990. Territorial salamanders evaluate size and
chitinous content of arthropod prey, pp. 111-126 in Hughes,
R.N. (editor). Behavouwrial Mechanisms of Food Selection.
Springer-Verlag, Heidelberg. 886 pp.
Jaeger, R.G. and Barnard, D.E. 1981. Foraging tactics of a
terrestrial salamander: Choice of diet in structurally simple
environments. American Naturalist 117: 639-664.
Pague, C.A. and Mitchell, J.C. 1991. Plethodon hubrichti Thurow,
Peaks of Otter Salamander, pp. 436-437 in Terwilliger, K.
(coordinator). Virginia’s Endangered Species. McDonald and
Woodward Publication Co., Blacksburg, Virgina. 672 pp.
Petranka, J.W., Brannon, M.P, Hopey, M.E., and Smith, C.K.
1994. Effects of timber harvesting on low elevation popula-
tions of southern Appalachian salamanders. Forest Ecology
and Management 67: 135-147.
Petranka, J.W., Eldridge, M.E., and Haley, K.E. 1993.
Effects of timber harvesting on southern Appalachian
salamanders. Conservation Biology 7: 363-370.
Pough, F.H., Smith, E.M., Rhodes, D.H., and Callazo, A.
1987. The abundance of salamanders in forest stands with
different histories of disturbance. Forest Ecology and
Management 20: 1-9.
Wicknick, J.A. 1995. Interspecific competition and territoriality
between a widespread species of salamander and a species with
alimited range. Ph.D. Dissertation, University of Southwestern
Louisiana, Lafayatte, Lowisiana. 152 pp.
Joseph C. Mitchell received his Ph.D. from the University
of Tennessee in 1982. Presently, he is an Adjunct Professor
in the School of Continuing Studies at the University of
Richmond and self-employed. Dr. Mitchell has published
over 100 papers and magazine articles, as well as two books.
Dr. Mitchell was the past secretary for the Herpetologists’
League and is now the president-elect.
Jill A. Wicknick received her Ph.D. from the University
of Southwestern Louisiana in 1995. She also holds a
postdoctoral degree from the same institution. Dr: Wicknick
specializes in the behavioral ecology of salamanders. Dr:
Wicknick has published in herpetological journals and the
highly respected journals Animal Behavior and Ecology.
Carl D. Anthony recewed his Ph.D. from the University
of Southwestern Louisiana in 1995. Dr. Anthony is finish-
ing his postdoctoral appointment at USWLA under Robert
Jaeger and will serve as an Assistant Professor at John
Carroll University starting Fall 1996. Carl specializes in
salamander behavioral ecology and parasitology. Dr. An-
thony has published in herpetological journals as well as
the journal Ecology.
ee
© Amphibian & Reptile Conservation
Printed in the United States. All rights reserved
Amphibian & Reptile Conservation (Fall 1996), 1(1): 20-23
Population Biology and Herpetological
Conservation: A Cautionary Note
Andrew Storfer
Center for Ecology, Evolution and Behavior, 101 TH. Morgan Hall, University of Kentucky, Lexington,
Kentucky 40506-0225
Abstract
Common endangered species conservation management practices, such as translocations and captive breed-
ing and release of individuals, tend to mix populations. This mixture is most often assumed to be benefi-
cial because it increases effective population size and may also increase genetic variation. However, when
populations are adapted to different local conditions, mixing populations can dilute local adaptation and
could theoretically result in population declines. Resource managers should therefore be careful when the
mixture of populations is considered, particularly in low dispersing species that are prone to local adap-
tation, such as salamanders.
Key Words
Demographics, gene flow, conservation, populations, management, inbreeding depression, amphibian, rep-
tile
Major concerns for small population
management
There are two major concerns in the conservation
of small populations: genetic and demographic fac-
tors. The genetic factors are inbreeding depression
and the loss of genetic variation through genetic
drift. Inbreeding depression threatens and reduc-
ing small populations with the possibility of unmask-
ing recessive deleterious alleles and a reduction in
heterozygosity. Each of these factors can, in turn,
affect individual fecundity and consequently reduce
population growth. When genetic variation is lost,
a population may decline because of its potential in-
ability to track environmental shifts (Frankel and
Soulé 1981; Lande and Barrowclough 1987). Demo-
graphic factors, such as the allee affect, where a
population is so small that individuals have difficulty
finding mates, can also severely affect small popu-
JAMIE K. REASER
COMMENTARY (Scientific)
lations. These factors are becoming more widely
recognized as tantamount to species extinction’s
(Lande 1988; Schemske, et al. 1994).
There is some debate about whether genetic
or demographic factors should be the primary focus
in the study and management of small populations.
However, the conservation programs that deal spe-
cifically with threatened or endangered populations
are often solutions to both problems. These manage-
ment schemes include: movement corridors between
nature reserves, captive breeding and release of in-
dividuals, and translocations of individuals between
populations. All of these strategies tend to enhance
gene flow, or the movement of individuals and the
integration of their genes, among populations.
The issue of Gene Flow
All of these potential solutions, therefore, are based
upon the assumption that gene flow is beneficial to
managed populations. Population genetic theory in-
dicates that gene flow can act as a “creative force”
by maintaining genetic variation and increasing ef-
fective population size and thus combating the nega-
tive consequences of inbreeding depression and de-
mographic stochasticity (Slatkin 1987). However,
there has been little consideration of the potential
detrimental effects that gene flow may have on popu-
lations despite the theory and empirical data to sup-
port such effects. For example, high levels of gene
flow among populations with different environments
(and consequently different selection regimes) can
swamp local adaptation and drive populations to
adapt to the average of local conditions (Wright
1951; Slatkin 1987); therefore, populations are not
well adapted. Associated with this swamping of lo-
cal adaptation can be a reduction in fitness, which is
known as outbreeding depression (Templeton 1986).
In the long-term, gene flow can also preclude sub-
population differentiation and eventually prevent
speciation by maintaining genetic contact (Mayr
1963, 1969).
Along with many examples of the enhancement
of gene flow resulting in the decline of managed popu-
lations (see Greig 1979), there are also examples of
gene flow acting as an evolutionary constraint for
natural amphibian and reptile species. A well-known
reptilian example is the Lake Erie water snake,
Nerodia sipedon. Banded snakes were found to be
more cryptic (and much more common) in the wooded
areas of the mainland surrounding Lake Erie, and
Correspondence, tel: (606) 323-5491, fax: (606) 257-1717, e-mail:
Storf@ceeb.uky.edu
A Cautionary Note on Herpetological Conservation 21
unbanded snakes were more common and cryptic in
the open, rocky areas of the islands (Camin and
Ehrlich 1958; King 1992). Varying degrees of band-
ing have consistently been found in the juvenile is-
land populations, despite its selective disadvantage
(Camin and Ehrlich 1958; King 1987). Observations
of snake dispersal and recent estimates of gene flow
indicated that individuals were dispersing from the
mainland to the islands (King 1987). Quantitative
genetic analyses indicated that color pattern was
largely genetically determined and that selection, in
the absence of gene flow, was strong enough to elimi-
nate the noneryptic (or banded) morph on the islands
(King 1993a). King recently concluded that gene flow
from the mainland population was overwhelming the
effect of selection, which caused an observed decline
in population fitness (King 1987, 1993a, 19930).
One amphibian example comes from research on
the stream-breeding salamander, Ambystoma
barbowri. In some streams that are deep enough to
support permanent pools, A. barbowri larvae face a
major selection pressure that comes from predatory
green sunfish, Lepomis cyanellis. In these streams the
optimal larval strategy entails reduced activity level
to remain inconspicuous and avoid predation (Sih, et al.
1988; Kats, et al. 1988). In other shallow streams, a key
selection pressure is habitat ephemerality, where the
optimal larval strategy entails increased activity level
to feed rapidly in order to reach a large enough size to
metamorphose before the stream dries up (Petranka
and Sih 1987). Recent work has shown that gene flow
is high enough between these two population types to
swamp local adaptation (Storfer, wnpublished data).
Additional data indicate that this gene flow has caused
adaptation to the average of local conditions because
a number of behavioral (7.e., refuge use, escape re-
sponse) and life history assays (7.e., stage at hatching)
associated with predator avoidance have been shown
not to differ significantly between two populations (one
with fish, the other ephemeral) connected by gene flow
(Storfer, wnpublished data). Even so, isolated popula-
tions do differ significantly in those traits associated
with fish avoidance. Therefore, gene flow may be
swamping local adaptation in some populations of A.
barbown and making them potentially more suscep-
tible to fish predation.
Management implications
Gene flow can play a major role in the management
of threatened or endangered species. In particular,
it is important for conservation biologists and re-
source managers to gain a better understanding of
gene flow as a constraining force. That is, there is
22 Andrew Storfer
enough uncertainty about whether gene flow acts as
a disrupter of local adaptation to warrant caution
toward management plans that include the enhance-
ment of interpopulation connection and gene flow.
Gene flow can be a particularly important issue for
amphibian species. For example, salamanders have
typically low levels of gene flow, thus making popu-
lations naturally subdivided (Larsen et al. 1984;
Slatkin 1985). Enhancement of gene flow for conser-
vation purposes may therefore not be warranted
based upon the fact that natural historical associa-
tions may not exist between the populations. It is
therefore important to conduct more studies of gene
flow in amphibian and reptile species, especially
since techniques of measuring gene flow have be-
come more and more accessible (see Slatkin 1985).
Studies of gene flow provide insight into population
structure and historical associations between popu-
lations helping resource managers to avoid mixture
of populations without high levels of gene flow, ex-
hibited by many amphibians species in nature.
It is essential to conduct ecological surveys of
habitat types and major selection pressures that af-
fect species which are management candidates, such
as limiting resources (7.e., prey, space, nest sites, etc.),
major predators, and intrinsic habitat differences
(7.e., climatic differences). These brief surveys would
not slow critical conservation decisions, yet they
would provide extremely valuable information so
that populations that are most ecologically alike
could be those that are mixed (when such manage-
ment may be warranted). Without such data, re-
source managers cannot be sure that mixing popula-
tions will not negatively affect the very populations
that they are trying to save.
References
Camin, J. and Ehrlich, P 1958. Natural selection in water snakes,
(Natria sipedon L.) on islands in Lake Erie. Evolution 12: 504-511.
Frankel, O.H. and Soulé, M.E. 1981. Conservation and Evolution.
Cambridge University Press. Cambridge, United Kingdom.
327 pp.
Greig, J.C. 1979. Principles of genetic conservation in relation to
wildlife management in South Africa. South African Journal of
Wildlife Restoration 9: 57-78.
Kats, L.B., Petrauka, J.W., and Sih, A. 1988. Antipredator
responses and the persistence of amphibian larvae with fishes.
Ecology 69: 1865-1870.
King, R.B. 1993a. Color pattern variation in lake Erie water
snakes: Prediction and measurement of natural selection.
Evolution 47: 1819-1833.
King, R.B. 1993b. Color pattern in lake Erie water snakes:
Inheritance. Canadian Journal of Zoology 71: 1985-1990.
COMMENTARY (Scientific)
King, R.B. 1992. Lake Erie water snakes revisited: Morph
and age specific variation in relative crypsis. Evolutionary
Ecology 6: 115-124.
King, R.B. 1987. Color pattern polymorphism in the lake Erie
water snake, Nerodia sipedon insularium. Evolution 41:
241-255.
Lande, R. 1988. Genetics and demography in biological con-
servation. Science 241: 1455-1460.
Lande, R. and Barrowclough, G.F. 1987. Effective population
size, genetic variation, and their use in population manage-
ment, pp. 87-125 in Soulé, M.E. (editor). Viable Populations
for Conservation. Cambridge University Press, Cam-
bridge, United Kingdom. 189 pp.
Larsen, A., Wake, D.B., and Yanev, K.P. 1984. Measuring
gene flow among populations having high levels of genetic
fragmentation. Genetics 106: 293-308.
Mayr, E. 1963. Animal Species and Evolution. Harvard
University Press, Cambridge, Massachusetts. 797 pp.
Mayr, E. 1969. Principle of Systematic Zoology. McGraw-
Hill, New York, New York. 428 pp.
Petranka, J.W. and Sih, A. 1987. Habitat duration, length of the
larval period and the evolution of a complex life cycle of an
amphibian. Hvolution 41: 1347-1356.
Schemske, D. et al. 1994. Evaluating approaches to the
conservation of rare and endangered plants. Ecology 75: 584-
606.
Sih, A., Petranka, J.W., and Kats, L.B. 1988. The dynamics
of prey refuge use: a model and tests with sunfish and
salamander larvae. American Naturalist 132: 463-483.
Slatkin, M. 1985. Gene flow in natural populations. Annual
Review of Ecology and Systematics 16: 393-430.
Slatkin, M. 1987. Gene flow and the geographic structure of
natural populations. Science 236: 787-792.
Templeton, A.R. 1986. Coadaptation and outbreeding depres-
sion, pp. 105-116 in Soulé, M.E. (editor). Conservation
Biology: The Science of Scarcity and Diversity. Sinauer
Associates, Sunderland, Massachusetts. 584 pp.
Wright, S. 1951. The genetical structure of populations. Annals
of Eugenics 15: 323-354.
After a brief stint as a Fish and Wildlife Biologist for the
US Fish and Wildlife Service in 1992, Andrew Storfer began
work as a graduate student in the Center for Ecology,
Evolution and Behavior at the University of Kentucky.
Since that time, he has been studying the evolutionary effects
of gene flow in a stream-dwelling salamander that is endemic
to Kentucky. His research incorporates aspects of genetics,
population ecology and behavior in an unusually integrative
project. Andrew expects his Ph.D. in May of 1997.
ee
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© Amphibian & Reptile Conservation
Amphibian & Reptile Conservation (Fall 1996), 1(1): 24-26
Printed in the United States. All rights reserved
Herpetofauna in the Wildlife Trade and
Nature: On the Difficulty of Estimation
Allen Salzberg
New York Turtle and Tortoise Society, 163 Amsterdam Avenue, Suite 365, New York, New York 10023-5001
Key Words
Population estimation, numbers, wildlife trade, reptiles, amphibians
Abstract
Due to the compleaity of animal
populations both in the wild and
the marketplace, it is difficult for
scientists to estimate the actual
numbers. Obstacles in the mar-
ketplace include: illegal activities,
inefficient estimation procedures,
and understaffed monitoring
agencies. Most experts agree that
the existing numbers often times
are estimated extremely low. All
numbers should be used with cau-
tion. Further research and tighter
controls are called for.
One of the first things we learn as
children is the art of counting. As
adults, counting large numbers is
replaced by complicated estima-
tion techniques. Large numbers of
variables makes estimation diffi-
cult. Nowhere is this more true
than when trying to measure the
trade in reptiles and amphibians
and its impact on wild populations.
Simply, the problem is that
reliable numbers of how many
animals are being sold, collected,
or in the wild do not exist. At
least, numbers which are precise.
Take for example the
seemingly simple act of esti-
mating how many animals are
sold in a given year. Any well run
business has this statistic but is
Correspondence, e-mail: Asalzberg@
aol.com
reluctant to make it public un-
less it is somehow mandated by
law. This is especially true of
some dealers and collectors of
live animals who are afraid of a
sudden surge in a certain species
attracting the unwanted
attention of their competi-
tion, scientists, conserva-
tionists or law enforce-
ment officers (Enge 1994).
So ruling out the pass-
ing of new laws, for example,
how does one find out how
many reptiles and amphib-
ians are sold within the
United States? Currently
the only numbers on pet
herpetofauna in the U.S. that
exist are guesses made by
two recent pet industry sur-
veys (American Pet Prod-
ucts Manufacturers 1994,
American Veterinary Medi-
cal Association 1992). They
estimate in the United
States there are between
one to four million reptiles
and amphibians kept as pets.
Not only is this number
too broad to be usable, sur-
veys failed to ask specific
questions that would supply
data which would be more
useful. They asked pet owners if
they owned a snake or a turtle. If
they had asked, more specifically,
what species owners had then sci-
entists could better estimate how
many amphibians and reptiles are
caught and sold each year and
their impact on wild populations.
One would have better luck
trying to find the numbers of
herpetofauna being sold into and
out of the United States. At least
Greek tortoises from Jordan in their original
shipping crates. The shipper of these turtles had
a CITES permit for a specific number of animals.
But as many CITES shipments do, this shipment
had 43 more animals than allowed, so the excess
animals were confiscated. Due to infrequent
inspections most “padded” shipments go through,
indicating that official CITES numbers do not
reflect actual number traded.
there exists established trade data
bases; LEMIS, or The Law En-
forcement Management Informa-
tion System run by the U.S. Fish
& Wildlife Service and a count of
CITES animals traded, put out
© 1994 ANITA SALZBERG
COLUMN (Popular)
by England’s
World Conser-
vation Moni-
toring Centre
(WCMC).
LEMIS
shows that the
USS. either im-
ports or ex-
ports up to ten
million live or
dead reptiles
and amphib-
ians per year.
But even here
there are prob-
lems. LEMIS
doesn’t include
all species. The
all mysterious
category of non-CITES reptiles in-
cludes hundreds of thousands of
animals, of unknown species, each
year. Entire families of lizards
such as skinks, agamas and geckos
are only listed by their common
family name. And then certain rep-
tiles, like Graptemys and Apalone
are listed only by genus. (The situ-
ation for these two genera im-
proved last year with the addition
of several species for both). This
forced the wildlife officer entering
the data to list numerous species
and subspecies together, thus los-
ing valuable data. Documents are
most often wrong. The right thing
would be to open each crate and
count each animal, something
which few Fish & Wildlife Inspec-
tors have the time to do unless
they are suspicious of some wrong-
doing (Luiijf 1994).
The WCMC collects the
numbers submitted by CITES
countries of just CITES listed
animals which have been traded.
Though the species listing is more
exact, the problem here is that
WCMC has no enforcement or
checking ability. It’s purely an
honor system. So a local CITES
Baby sliders in a Florida dealer’s sho
6 million of these were shipped out of the US last year alone.
Estimation Difficulties in the Wildlife Trade and Nature 25
i 3
officer who is allowing a lot of
CITES animals out, and doesn’t
want to attract attention, might
just fudge his country’s numbers
(Groombridge 1994).
One common gap that both
these databases share is that
there is no record keeping of the
animals’ ages (Anderson 1994).
When it comes to reptiles, espe-
cially turtles, this is very impor-
tant information since collection
of adults does more harm to a wild
populations than of hatchlings
(Congdon 1994).
Counting is also a problem
when keeping track of wild popu-
lations. Examples exist of popula-
tion studies which have proven in-
accurate because they didn’t take
into account the animal’s natural
habits. This resulted in scientists
looking for animals where they
were least likely to find them (Gib-
bons 1993).
We also know that popula-
tions often naturally increase or
decrease tremendously, and only
long-term population studies take
that into account and more accu-
rately measure a population’s
trend (Gibbons 1993).
p awaiting shipment. It is estimated that over
Another
problem is who
does the col-
lecting. My fa-
vorite example
of this is a sto-
ry about Caret-
tochelys inculp-
ta. It originally
was believed
to be a very
rare species. It
turned out that
the scientists
who first dis-
covered this
animal were to-
tally dependent
on natives to
collect their
specimens. They asked the natives
to bring back any turtle they
caught when they would go
upriver. The problem was that the
natives like to eat Carettochelys
and so gave the scientists only the
bad tasting turtles (Pritchard 1993).
Now does this mean that
these numbers are useless? No.
Even if the numbers are incom-
plete they are useful. Most herpe-
tologists believe the numbers are
way too low, (Enge 1994) describ-
ing live animal shipments that are
smuggled through customs with
invoices listing the boxes’ con-
tents as light bulbs and so they
are never seen by Wildlife officers
(Luiijf 1993).
And like the canary in the
mines, LEMIS and WCMC data
can act as a warning to trade ac-
tivities, excessive or illegal.
Large numbers, even of just liz-
ards or softshells, suddenly being
reported can indicate to conserva-
tionists and scientists that further
study or action is necessary.
Recently, Togo and Ghana
was reporting to WCMC an in-
credible amount of reptiles being
exported, more than those small
© 1994 ALLEN SALZBERG
Se
26 Allen Salzberg
countries could possibly collect.
This tipped off wildlife officials to
investigate. They discovered that
Togo and Ghana was laundering
animals illegally collected from
other countries (Ventura 1994).
But still we have to figure
out how best to estimate amphib-
ians and reptiles, both collected
and traded (these two numbers
are usually different because ani-
mals do die in transit, or are kept
for personal use by the collector).
The U.S. Fish & Wildlife Service
needs to be encouraged to im-
prove LEMIS soit reflects the bio-
logical and natural history of the
animals being counted. Every
shipment should be opened and
counted. And any animal traded
should be on that system. Herpe-
tologists and conservationists
need to conduct their own surveys
of what is being sold in pet shops.
As for counting amphibians
oO
ia
int
5
x
a
z
a
4
3g
=
A typical storage set up for small snakes
at a Florida dealer. A common marketing
technique of animal dealers is to under-list
the amount of animals available in their
mail order catalogs—by as much as 90 %.
This creates the illusion of rarity and thus
boosts demand for the animal.
and reptiles in the wild, an excel-
lent reference is Measuring and
Monitoring Biological Diversity:
Standard Methods for Amphib-
ians (Heyer et al. 1994). Though I
look forward to an edition on rep-
tiles, this book, with its detailed
studies of different collection and
recording techniques, should give
anyone doing population studies
on reptiles help on how to formu-
late a useful, comparable, popula-
tion study.
Simply: You can’t save what
you don’t know you have.
References
Anderson, Mary. 1994. Personal Com-
munication. September. Independent
Researcher, Roanoke, Virginia.
American Pet Products Manufacturers
Association. 1994. 1994 National Pet
Owners Survey, pp. 180-197. American
Pet Products Manufacturers Asso-
ciation, Scarsdale, New York.
American Veterinary Medical Asso-
ciation. 1992. The Veterinary Service
Market for Companion Animals, 1992,
pp. 103-106. Center for Information
Management, American Veterinary Med-
ical Association, Scaumburg, Illinois.
120 pp.
Bright, E. 1994. An analysis of U.S. Fish
& Wildlife reptile and amphibian num-
bers in their LEMIS system. Unpub-
lished manuscript.
Conant, R. and Collins, J.T. 1991. A
Field Guide to Reptiles and Amphib-
ians Eastern/Central North America.
3rd Edition. Peterson Field Guide
Series. Houghton Mifflin Company,
Boston. 450 pp.
Congdon, J.D., Dunham, A.E., and Van
Loben Sels, R.C. 1993. Delayed sexual
maturity and demographics of Blanding’s
turtles (Emydoidea blandingii): Impli-
cations for conservation and manage-
ment on longlived organisms. Conser-
vation Biology (2): 387-399.
Enge, K.M. 1994. Herptile Use and
Trade in Florida. Florida Game and Fresh
Water Commission Nongame Wildlife
COLUMN (Popular)
Program Final Performance Report,
Tallahasse, Florida. 102 pp.
Gibbons, J.W. 1993. Measuring declines
and natural variation in populations of
turtles: spatial lessons from longterm
studies. Presentation at Conservation,
Restoration, and Management of
Tortoises and Turtles an International
Conference. July, Purchase, New York.
Groombridge, L. 1994. Personal Com-
munications. World Conservation
Monitoring Centre, Cambridge, United
Kingdom.
Heyer, W. R., Donnelly, M.A., MeDiar-
mid, R.W., Hayek, L.C., and Fraser,
M.S. 1994. Measuring and Monitoring
Biological Diversity: Standard Methods
for Amphibians. Smithsonian Institution
Press, Washington, DC.
Luiijf, W. 1993. The role of CITES in
controlling tortoise trade. Presentation at
Conservation, Restoration, Management
of Tortoises and Turtles and Interna-
tional Conference. July, Purchase, New
York.
Pritchard, Dr. PRC.H. 1993. Personal
Communications. Vice Preseident, Flor-
ida Audubon Society, Casseberry,
Florida.
Ventura, J. 1994. Personal Communica-
tion. Supervisory Wildlife Inspector,
U.S. Fish & Wildlife Service, Port of
New York, John F. Kennedy Airport.
Warwick, C. 1994. Personal Commun-
ications. People’s Trust for Endangered
Species, Surrey, England.
Allen Salzberg’s interests in environ-
mental issues began back in 1965 after
he saw his first wild turtle. Mr: Salzberg
has written numerous articles on the
environment for magazines including
Health, Outside, and The New York
Times. Recently, he completed a report
titled The Preliminary Report: Live
Freshwater Turtle & Tortoise Trade in
the United States for the US Humane
Society and Humane Society Interna-
tional. He is presently working on wp-
dating and expanding that report to
include all reptiles. With his wife, Anita
Baskin Salzberg, he has cowritten a
children’s book on turtles for “all those
kids like me back in 65.”
ee
NEWS and NOTES
ANNOUNCEMENTS
New Organization: African
Herpetofaunal Biodiversity Programme
(AHBP)
The AHBP is a new and developing
programme designed to identify, assess,
inventory, and monitor conservation areas
in Africa and Madagascar. Since herpe-
tofaunas’ remains much neglected with
only the crocodilians and, to a degree, the
chelonians receiving major attention rep-
tiles and amphibians will be a major focus
of the AHBP In order to proceed further,
itis necessary to elect a coordinating com-
mittee, find a home for the programme (at
present, Pietermaritzburg, South Africa),
and to prepare initiatives to secure start-
up funding and project linkages. Help is
also needed in setting up a permanent list
server site to promote communication
between collaborators, adding new mem-
bers to the administrating committee, and
assisting with other projects. All those in-
terested or who can help are urged to con-
tact Lynn Raw, PO Box 200, Merrivale,
3291, South Africa. Tel: +27-331-
460796, fax: +27-331-460895, e-mail:
Raw @zoology.unp.ac.za. Presently, e-mail
and a mailing list (AFRIHERP-L) are
being used to aid communication between
AHBP members. To subscribe to the mail-
ing list send this e-mail message
(subscribe AFRIHERP-L “Your name”)
to: listproc@wemc.org.uk. To post mes-
sages to the list send them to:
Afriherp@wemce.org.uk. The list adminis-
trator is Lynn Raw. The AHBP “Interim”
Committee members are: Lynn Raw
(South Africa), Chairman (coordinating
facilitator); Ivan Ineich (France), Secre-
tary; Craig Hassapakis (United States),
Public Relations Officer; Preston
Hardison (United States), Data Manage-
ment Officer; Neil Burgess (Denmark),
Mapping; Michael Lambert (United King-
dom), Project Coordinator, and Martin
Kundrat (Slovak Republic) Project Coor-
dinator.
Dr. lan R. Swingland named Professor
of Conservation Biology
Dr. Ian R. Swingland has been named
Professor and Chair of Conservation Bi-
ology at the Durrell Institute of Conser-
vation and Ecology (DICE), University of
Kent, Canterbury, England. Following
many years of field work on Aldabra gi-
ant tortoises, Dr. Swingland was also the
Director of the First World Congress of
Herpetology, held in Canterbury in 1989,
instrumental in the formation of the
IUCN/SSC Tortoise and Freshwater
Turtle Specialist group, and has worked
for the past six years on the development
of DICE. DICE now has an international
program of postgraduate training in con-
servation biology, drawing students from
around the world for a unique Masters of
Science (M.Sc.) degree in Conservation
Biology.
MEETINGS
Third World Congress of Herpetology,
2-10 August, 1997
For more information contact: c/o Czech
Medical Association, J.E. Purkyne, Con-
gress Department, PO Box 88, Sokolska
31, 120 26, Praha 2, Czech Republic. Tel:
(++42-2) 296889 (or) (+ +42-2) 249151195,
fax: (++42-2) 294610 (or) (++42-2)
24216836. The web site address is: http://
www.gli.cas.cz/herpet/
Conference to be Hosted in Vietnam:
Conservation and Biodiversity of
Amphibians and Reptiles of Tropical
Rain Forests
The proposed site for the conference is
Hanoi, Vietnam, sometime in 1998. Those
interested should contact one of the fol-
lowing: Dr. Harold Heatwole, Department
of Zoology, Box 7617, North Carolina
State University, Raleigh, North Carolina
27695 USA. Tel: (919) 515-2741, fax: (919)
515-5327; Dr. Natalia Ananyeva, Depart-
ment of Herpetology, Zoological Institute,
St. Petersburg 199034, Russia. E-mail:
any@zisp.spb.su; Dr. Cao Van Sung, Insti-
tute of Ecology and Biological Resources,
Nghia Do, Tu Liem, Hanoi, Vietnam. Tel:
361 440, fax: (844) 361 196, e-mail:
sung@iebr.ac.vn
CALL FOR INFORMATION
Live Reptile and Amphibian Trade
Database
Information for a database for analysis is
being gathered on the trade in live rep-
tiles and amphibians. The purpose of the
database is to monitor the live trade and
issue occasional reports and suggestions
27
on how to improve the trade, making it
more humane and environmentally
sound (one recent use was to get the
North American box turtle listed as an
CITES II species). Notes of anecdotes,
stories of people handling herpetofauna
for the live animal trade would be use-
ful additions to the database. These in-
clude copies of articles or papers on the
trade’s effect on wild populations, sto-
ries of arrests, or visits to turtle farms.
Include other topics which are relevant,
whether favorable for the trade or not.
Farming of amphibians and reptiles and
sustainable harvesting stories are also
appreciated, as well as suggestions of
how to improve the trade. Send infor-
mation to: Allen Salzberg, 6787 Booth
Street 5B, Forest Hills, New York
11375. Tel/fax: (718) 275-3307, e-mail:
Asalzberg@aol.com
BOOKS RECEIVED
Murphy, J.B., Adler, K., and Collins, J.T. (editors). 1994.
Captive Management and Conservation of Amphibians
and Reptiles. Society for the Study of Amphibians and
Reptiles, Ithaca, New York. 408 pp.
Musgrave, R.S. and Mary A.S. 1993. State Wildlife Laws
Handbook. Government Institutes, Inc., Rockville,
Maryland. 840 pp.
Cogger, H.G., Cameron, E.E., Sadlier, R., and Eggler, P
1993. The Action Plan for Australian Reptiles. Aus-
tralian Nature Conservation Agency, Canberra, ACT. 254
pp.
Shine, R. 1991. Australian Snakes: A Natural History.
Cornell University Press, Ithaca, New York. 223 pp.
Roughgarden, J. 1995. Anolis Lizards of the Caribbean:
Ecology, Evolution, and Plate Tectonics. Oxford Uni-
versity Press, New York. 200 pp.
Stebbins, R.C. and Cohen, N.W. 1995.A Natural History
of Amphibians. Princeton University Press, Princeton,
New Jersey. 316 pp.
Hudson, R., Alberts, A., Ellis, S., and Byers, O. 1994. Con-
servation Assessment and Management Plan for
Iguanidae and Varanidae. IUCN/SSC Conservation
Breeding Specialist Group, Apple Valley, Minnesota.
IUDZG/CBSG (IUCN/SSC). 1993. The World Zoo Con-
servation Strategy: The Role of the Zoos and Aquaria
of the World in Global Conservation. Chicago Zoologi-
cal Society, Brookfield, Illinois. 76 pp.
Wiese, R.J. and Hutchins, M. 1994. Species Survival
Plans: Strategies for Wildlife Conservation. Ameri-
can Zoo and Aquarium Association, Wheeling, West Vir-
ginia. 64 pp.
Slavens, F. and Slavens, K. 1994. Reptiles and Amphib-
ians in Captivity, Breeding-Longevity and Inventory,
Current January 1, 1994. Slaveware, Seattle, Washing-
ton. 532 pp.
Hunter, Jr., M.L., Albright, J., and Arbuckle, J. (editors).
1992. The Amphibians and Reptiles of Maine. Maine Ag-
ricultural Experimental Station Bulletin 838. 188 pp.
28
EDITORIAL
LITERATURE
Cree, A., et al. 1995. Reproduction of a rare New Zealand reptile, the tuatara Sphe-
nodon punctatus, on rat-free and rat-inhabited islands. Conservation Biology 9(2):
373-383.
Kiesecker, J.M. and Blaustein, A.R. 1995. Synergism between UV-B radiation and a
pathogen magnifies amphibian embryo mortality in nature. Proceedings of the National
Academy of Sciences (USA) 92(24): 11049-11052.
Durbin, J., Rajafetra, V, Reid, D., and Razandrizanakanirina, D. 1996. Local people and
project Angonoka—conservation of the ploughshare tortoise in north-western Mada-
gascar. Oryx 30(2): 113-120.
Kaiser, H. 1994 The conservation status of Lesser Antillean frogs. Herpetological
Natural History 2(2): 41-56,
Daugherty, C.H., Patterson, G.B., and Hitehmough, R.A. 1994. Taxonomic and conser-
vation review of New Zealand herpetofauna. New Zealand Journal of Zoology 21(4):
317-323.
Brown, D. 1994. Transfer of Hamilton’s frog, Leiopelma hamiltoni, to a newly cre-
ated habitat on Stephen’s Island, New Zealand. New Zealand Journal of Zoology 21(4):
425-430.
Towns, D.R. 1994. The role of ecological restoration in the conservation of Whitaker’s
skink (Cyclodina whitakeri), arare New Zealand lizard (Lacertilia: Scincidae). New
Zealand Journal of Zoology 21(4): 457-471.
Herpetological Conservation. This new publication is a book-length monograph se-
ries on the conservation of amphibians and reptiles. HC is published approximately once
per year and will concentrate on a single subject. For information contact Paul Stephen
Corn, Editor, National Biological Service, Aldo Leopold Wilderness Research Insti-
tute, 709 East Beckwith Avenue, PO Box 8089, Missoula, Montana USA. Tel: (406) 542-
4190, fax: (406) 543-2663, e-mail: Steve_Corn@nbs.gov.
ce
ACKNOWLEDGMENTS
A great tribute must be given to the early support I received for the journal from sub-
scribers, editors, advisors, and many others. Of great support were Jamie K. Reaser,
David Galbraith, Andrew T. Holyeross, Bill Love, Kraig Adler, Ken Dodd, Russell
Mittermeier, Michael Hutchins, Aaron Bauer, Allison Alberts, Jonathan Ballou, Joseph
T. Collins, Carl Gans, Howard Lawler, Roy McDiarmid, George B. Rabb, Hobart Smith,
Lee Fitzgerald, Steven Garber, Julian Lee, Joe Mitchell, Henry Mushinsky, Christo-
pher Raxworthy, Nelson da Silva, Andrew Storfer, Robert Wiese, Frank Slavens, Michael
and Patrica Fogden, Anders Rhodin, Karen Toepfer, Robert Hansen, John Baker, Eric
Thiss, Lynn Raw, Martin Kundrat, Allen Salzberg, Dave Adams, Amy Stout, Sam Ashely,
Dave Owens, Ian Straw, Bret Bottger, and early subscribers, too many to list here.
Needed information and help have always been provided by the excellent staffs at the
IUCN/SSC Conservation Breeding Specialist Group (CBSG), International Species
Inventory System (ISIS), and World Conservation Monitoring Centre (WCMC). A
special thanks is extended to Cornell University Press, Princeton University Press,
and American Graphics for all their help. Helpful assistance was provided by Brigham
Young University particularly the Zoology Department, Computer Services, the Monte
L. Bean Life Science Museum, the Harold B. Lee Library (HBLL), and the HBLL
Interlibrary Loan Office. Individuals from Brigham Young University I would like to
personally thank for there help and encouragement are Richard Tolman, Hal Black,
Clayton White, Jack Sites, Dr. Shiozawa, Nathan Smith, Wilmer Tanner, Doug Cox, Ri-
chard Jensen, Richard Soares, Roger Flick, John Higginbotham, and Stan Peters. Spe-
cial thanks goes to Walter “Wally” Tordoff, III at California State University at
Stanislaus for help when I was a young bubbling herpetologist. A tribute is given in honor
to the late Michael Parks of Ceres High School, Ceres, California. His friendship, humor,
and enthusiasm for education and the sciences will always be remembered. I would also
like to acknowledge some friendships which have always been there when I needed them:
Scott Pittel, Mark Adkison, Dave Pennock, Sally Midgley, Michael Jensen, M.D., Troy
Rowan, David Owens, Richard Soares and Tony Marren. The idea to publish a journal
and technical support was provided by Jay Vilhena. Without Jay I would not be publish-
ing this journal. At a very critical time the project would have never been completed
without the help, support, and friendship of Carin Young. Special thanks go to the fol-
lowing individuals who have exhibited a special effort to myself individually and in this
project: Jamie K. Reaser, Bob Wiese, Jay Vilhena, Troy Rowan, David Owens, Tony
Marren, Richard Soares, and Carin Young. Last but not least I would like to acknowl-
edge my parents Anthony and Frances Hassapakis and my two brothers Steve and Greg
Hassapakis. Thank you all for your unfailing support and friendship.
EDITORIAL
Finally, I am proud to announce the first issue of Amphibian & Rep-
tile Conservation!!! The work has been under construction now for
over 1 1/2 years with many improvements and refinements being
added. It is my hope that this publication will become a valuable
tool for conservation and in educating people about the need to pre-
serve our herptofauna heritage.
It was my vision to publish a journal about amphibian and reptile con-
servation that would appeal to as many people as possible while consist-
ing of the best science available. In the past this would have meant pub-
lishing a science journal with limited appeal in the private sector. I have
always believed that both could be accomplished together, a readable
science journal that the public could better understand and enjoy. I think
Ihave accomplished this vision by creating a journal style with elements
of a magazine (color photos and pleasing graphic design), scientific infor-
mation (original data, scientific article format, and respected science au-
thors), and a newsletter type section for all other useful information. The
result will be a stronger more educated public who have more time, money,
and enthusiasm to contribute to conservation; something that is often in
short supply in the conservation community. The professional scientific
conservation community is a limited resource and is often strained to its
limits. Conservation is too important of an area to limit to these hard
working individuals. There is power in numbers. By increasing our read-
ership and their education we will have a greater impact toward helping
others do something about the conservation efforts of amphibians and rep-
tiles. Let’s not take a back seat to our commitment to educating ourselves
and others about what is being done in the field of herpetological conser-
vation. Make a commitment now to become a subscriber to ARC and con-
tribute by writing articles, reporting your research finds, loaning your
prized photographs, and telling others of this exciting new venue for con-
servation. What better way is there than this to show your commitment
and support for reptile and amphibian conservation?
All of the good will and individual effort in the world will not make
ARC asuccess without your help, support, and valuable feedback. What
is being done in your part of the world to conserve its herpetofauna? Why
not drop ARC a line and tell us. Now with computers and the Internet
ARCis only an e-mail away!
It is my hope that through Amphibian & Reptile Conservation we can
all unite and make a big difference toward the conservation efforts of am-
phibians and reptiles worldwide, as well as all life.
With sincere hopes,
Craig Hassapakis
DEDICATION
This journal is dedicated in honor of my grandparents, Alla Mae “Mona” and
Lawton Lail “Grandad” Hendricks. Our rides through the countryside, their spe-
cial friendship and unfailing support for all my projects will never be forgotten.
Lawton Lail “Grandad” Alla Mae “Mona”
Hendricks Hendricks
(1911 - 1976) (1914- )
MT. HOOD, OREGON, USA (1941)
AMPHIBIAN & REPTILE
CONSERVATION
THE INTERNATIONAL JOURNAL DEVOTED TO THE WORLDWIDE PRESERVATION
AND MANAGEMENT OF AMPHIBIAN AND REPTILIAN DIVERSITY
Scope
Amphibian & Reptile Conservation (ARC) is an international peer reviewed journal
devoted to the worldwide preservation and management of amphibian and reptilian
diversity.
Goals
The primary goals of Amphibian & Reptile Conservation are to:
+ Provide a forum for scientists and others to publish data pertinent to amphib-
ian and reptile conservation.
+ Report on herpetological conservation efforts worldwide.
+ Assure the protection of all taxa of amphibians and reptiles, particularly those
= - Seon, need ete
on the’ ‘numbers of animals \ Gndividuals, populations, and species) in
“captivity, the witdlife Aili he tS
maatiee for S abiciion which inform, exe Feport new iscoyeries, offer: solutions,
present new information, and summarize rese ch) mpiemaand areas of interest.
Manuscripts may cover any aspect of amphibian an biology with an emphasis.
on conservation. ARC is particularly interested in ma aeene dealing with topical re-
views, zoo biology, population status (both in captivity and nature), rare, threatened,
and endangered species, human exploitation, resource management, , geographic dis-
tribution, herpetofaunal diversity, area checklists, exploration and discovery, allareas
of conservation biology (from such topics as economics and politics to held biology and .
techniques) and articles which relate animals to specific geographical locations and
countries (¢.g., The Endangered Frogs of Costa Rica and The Lizard Fatigaigitlads>
gascar with Particular Emphasis on Threatened Species). 3 {
INVITATION TO PARTICIPATE ON EDITORIAL AND
ADVISORY BOARDS
Amphibian & Reptile Conservation is interested in attracting qualified individuals to
participate on the editorial review or advisory boards. Please send Curriculum Vita,
list of publications, and a list of other offices and/or positions held to the editor for
consideration at 2255 North University Parkway No. 15, Provo, Utah 84604-7506 USA.
WRITER'S GUIDELINES AND MANUSCRIPT
PREPARATION
General Information
Amphibian & Reptile Conservation is intended for a wide readership from the ama-
teur to the professional. The study of conservation addresses a tremendous variety
of disciplines and occupations, experts in one area may not be knowledgeable in others.
Contributors are therefore encouraged to present their subject matter in simple, lu-
cid, and concise terms, appealing to amateur herpetologists, the general public, and sci-
entists. Articles should include abstracts, explanatory text (especially statistical and
mathematical methodologies), full references, ample photographs, tables, figures, dia-
grams, and other illustrations to help elucidate the text, whenever possible. All contri-
butions should be submitted to Amphibian & Reptile Conservation exclusively. If
accepted, papers and all other contributions become copyright of Amphibian & Rep-
tile Conservation. Full horizontal journal size measures 8 x 10.75 inches.
Comments
Comments are invited to ensure that Amphibian & Reptile Conservation is the best
journal possible. Feedback is very important. We appreciate submission of comments,
suggestions for improvements and/or corrections.
Publication Categories
All contributions should be designed to one of the following categories (see below
Feature and Scientific Articles, Departments [Short Communications, Commentar-
ies, Book Reviews, News and Notes - Brief Reports, Announcements, Meetings, Calls
for Information, Books Received, Literature, and Forthcoming], Editorial, and Col-
umn) for publication. However, new sections to the journal are encouraged. Proposals
should be discussed with the editor directly.
FEATURE AND SCIENTIFIC ARTICLES
Feature and Scientific Articles and other submissions listed in the Departments sec-
tion should follow the submission format listed below, unless otherwise stated. Fea-
Writer’s Guidelines and Manuscript Preparation 29
ture articles maybe popular science articles and reviews. Scientific articles include sci-
entific contributions which report new information based on original research and
reviews summarizing areas of research. Please label contributions as Feature, Sci-
entific, Review, Commentary, Popular or combination (7.e., Review [Popular]) in the
upper left hand corner of the title page. All articles should also contain the following
information and conform to the format below.
Title Page
The title page of the manuscript should consist of a single page and should include the
complete title of the paper; the names of the authors and their affiliations, address (in-
clude extra 4 digit zip code e.g., 94305-1901), fax and telephone numbers as well as e-
mail addresses (when available. If no e-mail is available please indicate by stating “No
e-mail available”); a short title (no more than 40 characters including spaces) to be used
as arunning header on top of nontitled pages; and the name of the person to whom edi-
torial correspondence, page proofs, and reprints requests should be sent. Please write
the total word count of the article in the upper right hand corner.
Abstract
A brief abstract (generally not exceeding 5% of the length of the text), intelligible without
reference to the main body of the text, should be provided to appear at the head of the
paper. At least six key words and/or phrases (to be used for subject matter indexing)
should appear on a separate line directly following the abstract.
Table of Contents
Provide a table of contents.
Manuscripts
Authors submitting manuscripts should send three paper copies, double-spaced, and
on one side of standard letter-sized paper, 21.5 x 28 centimeters (8.5 x 11 inches).
Manuscripts : should) peaubmitted i in electronic media and postal mailed along with the
the “Internet” (on Internet Submissions).
i satus of Amphibian & rele. Conservation for examples of style. For
further as tance on the proper ‘style and formatof manuscripts (not described in these
- guidelines), consult Scientific: Style and Format: Dhe CBE Manual for Authors, Edi-
tors, and. Publishers (6th edition, 1994, Council of ‘Biology Editors, Inc., 11 South
_-LaSalle, Suite’ 1400, ‘Chicago, ‘linols ss} Welk *
Submissions — Es
___ Final versions sof all! submissions to the Foumnal Brent be on3.1/2 Siac computer disks
or Iomega Zip disks suitable for IBM compatible computers. Please label all disks with
_ the name of the Software (e.g., Microsoft Word 6.0). ASCII is preferred but MicroSoft
Wor
“Submission of electronicillustrations i is strongly encouraged, but not required (see Te
lustrations, Bap All authors receive one copy of the journal free. ~~
“BE
Internet Submissions i y
The Internet provi¢ es.the, uickest method for subitting’ Simian PARC. es: ;
pecially near deadlines. This i is also the preferred method ito communicate direetly with
the editor, submit pr: oposals, and quickly? review. manuscripts for possible publication.
Pictures can also be retrieved over the Internet as as rencoded files.) , These are then de-
coded by the editor for reviewing, Detoder/encoder programs Ge WINCODE) can
be downloaded to your computeras shareware (free of charge) via | the Internet. Please
contact a computer consultant-or the. editor for specific details on-how'to s send and
retrieve encoded internet messages: eg mnder, what conditions to make Sbgigrrange-
ments.
Text ’
Regular articles should not exceed 10 manuscript} pages. RéviewScan be up to 20 pages =
in length and should include an introduction and conclusions:
will be considered, however,’ and the editor should be contacted to discuss s possible.
publication, for articles longer than the above standards. 3) Submission ions less fhan 4 pages >
are categorized as aShort Communication. The News and ote setin should only
include essential.in formation i in a succinet: fashion usually less,than nena Resiilts
and information should be presented i in clear format using photographs.
ustrations,
and tables to enhance the text whenever possible. A statement of compliance with the
guidelines for the use of animals in research, as published in Animal Behavior, Volume
43, 1992, must be provided in appropriate cases. For field research consult Pisani, G.R.
et al. 1987. Guidelines for the Use of Live Amphibians and Reptiles in Field Re-
search published by the Society for the Study of Amphibians and Reptiles. Nonstand-
ard abbreviations should be kept to a minimum and defined in the text. Vernacular names
can be used where appropriate, with the scientific name given in italics following the
common name of the first mention of the species. All measurements should be spelled
out on the first usage, with the abbreviation in parentheses, and the abbreviated form
used thereafter. Use the metric system unless English measurements are clearly more
appropriate (then give metric equivalents in parentheses). Indicate the approximate
placement of figures, tables, photographs, and maps on the manuscript. Formatting
such as bolding should be kept to a minimum and, if done, follow examples from the jour-
nal. Double-space lines between sentences. USE ONLY ONE SPACE AFTER ALL
PUNCTUATION MARKS SUCH AS PERIODS AND SEMICOLONS. DO NOT
hyphenate on line breaks (e.g., DO NOT justify the margins). Footnotes should be
avoided. Use italics for et al., i.e., ande.g. and scientific names of species. Use the word
“herpetofauna” in place of “herps.” Careful attention should be paid to accents and dia-
critical marks on foreign words. Add them by hand if not available on the typewriter
or computer.
‘and Word Perfect/are also aceeptable formats to submit manuscripts. Authors
__shétild:feeltree | to.suggest at least one referee qualified to judge the work objectively. \ ~
ortsJonger than'20 pages ~
+
30 Writer’s Guidelines and Manuscript Preparation
References
References should follow the Harvard system. When more than two authors are ref-
erenced in the text et al. follows the name of the first author, for example (Mittermeier
et al. 1996). Use initials for all names in the references except surnames leaving no
spaces between letters. Italicize book and journal titles using the language and spelling
of the original and give English translations, in square parentheses when needed. With
hard to obtain papers listed in the references, include contact information for obtain-
ing copies. List total page numbers for books at the end of the reference. For examples
of the format used see below.
Lande, R. and Barrowclough, G.H. 1987. Effective population size, genetic varia-
tion, and their use in population management, pp. 87-125 in Soulé, M_E. (editor).
Viable Populations for Conservation. Cambridge University Press, Cam-
bridge, United Kingdom. 189 pp.
Mittermeier, R.A., Werner, T.B., and Lees, A. 1996. New Caledonia—a conser-
vation imperative for an ancient land. Oryx 30(2): 104-112.
Stebbins, R.C. and Cohen, N.W. 1995. A Natural History of Amphibians.
Princeton University Press, Princeton, New Jersey. 316 pp.
Acknowledgments
Limit acknowledgments to individuals and organizations who contributed directly to
researching and funding. Keep them as brief as is consistent with courtesy.
Author Biography
A brief biography of the author is included at the end of each article. Information pro-
vided should be limited to titles, academic positions, employment, past and recent re-
search projects, and details containing brief descriptions of the authors and the over-
all research projects within which the published work has been carried out. This will
provide readers with an outline of the structure and objectives of the research teams,
or groups responsible for the work.
Correspondence
Designate a correspondence author to whom all communications concerning the paper
should be directed. List the most up-to-date telephone and fax numbers along with an
e-mail address. Promptly notify the editor if any of these numbers should change before
publication.
Illustrations
Illustrations (line drawings, diagrams and other illustrations such as Photographs,
Tables, and Figures—see below) should be submitted as both electronic versions and
hard copy with author’s name and figure number attached. These works should be
suitable for 50% reduction without loss of clarity. Exceptions are photographs where
90% reduction is acceptable. Submission of electronic illustrations are highly encour-
aged, but are not an absolute requirement. Submit illustrations on computer disk as
separate files from the text in EPS, WMF or DXF format or as quality hard copies.
Always include a printed copy of all electronic illustrations along with descriptive cap-
tions. It is suggested that individuals submitting electronic work contact the editor in
advance for more detailed instructions and clarification of techniques. Disks will usu-
ally not be returned.
Photographs
Halftone photographs (both color and black and white) should be submitted as 35 mm
slides or larger (large format transparencies 7.e¢., 4 x 5 inches). If slides are not avail-
able glossy prints should be selected of a large size if possible, e.g., 8 x 10 inches. Details
about the photographs (place, time, date, and description) should accompany all pictures.
A hard copy description of the photographs should be typed consecutively on separate
sheets of paper and also sent to the editor on a computer disc. If identifiable human
subjects are used in photographs a signed letter of consent must be included.
Tables
Tables must be typed double-spaced without vertical rules and should not duplicate any
material in the text or illustrations. Tables should be tab delimited using such programs
as Microsoft Notepad, Word, or Excel, and submitted in electronic media. Provide all
tables with complete but brief headings. Type them on separate sheets of paper, num-
ber them consecutively within the text, and include them on a computer disk.
Figures
Type figure legends double-spaced on consecutively numbered separate pages and
include them on a computer disk.
Proofs
Proofs will be sent whenever possible but because of publishing deadlines this may not
always be possible and/or necessary. After acceptance, papers may be edited to en-
hance clarity, The senior author will be notified of significant changes in content or style.
Page Charges
Page charges of $35 per page will be assessed for publication costs for those who have
grants or institutional support, $10 per page for others. This fee will be waived if a state-
ment is signed indicating that neither there is no institutional support for publication
and/or that $10 per page cannot be paid. Charges will be collected by the publisher. An
author's inability to pay will in no way influence acceptance of the paper for publication.
Offprints
The publisher will supply the author with 25 free offprints. An offprint order form will
be included with the page proofs and authors may order further offprints. For pricing,
please contact the editor.
DEPARTMENTS
Not all sections of the Departments section will appear in each issue of Amphibian &
Reptile Conservation and new sections will be added as warranted.
Short Communications
Papers shorter than four pages should be prepared as Short Communications.
Commentaries
Commentaries are explanatory or illustrative, systematic series of comments or
opinions based on critical notes or observations.
Book Reviews
Book Reviews are critical evaluations on books. Publishers requesting that their book(s)
be reviewed should contact the editor. Whenever possible two copies should be sent with
a written request to review. Persons wanting to review books should contact the editor
with a proposal.
News and Notes
News and Notes is the newsletter section of the journal. Submissions should be con-
fined to timely news of interest to the herpetological conservation community. These
contributions will be placed in one of the following subsections.
Brief Reports
Brief Reports should be fashioned after the “Briefly Section” in the Journal of
the Fauna & Flora Preservation Society (London), Orya.
Announcements
Announcements are important timely reports of activities in the herpetological
conservation community and should be reported here.
Meetings
Meetings includes references to upcoming conferences, symposiums, and
workshops.
Calls for Information
Calls for Information is a service provided to researchers. It is intended to elicit
contributions from others and to further the effort.
Books Received
Books Received is a listing of books received for review . It will include full bib-
liographic data and (sometimes) brief comments.
Literature
Literature cites important works which directly (or indirectly) relate to the
conservation of amphibians and reptiles. Recent literature is most often listed
but other important literature such as classical works may be also included.
Forthcoming
Forthcoming brings attention to future articles solicited or in press in Amphib-
ian & Reptile Conservation.
EDITORIAL
The editorial is a topical subject commentary which is usually written by the editor, On
oceasion, guest editorials may be solicited.
COLUMN
Columns are assigned by the editor. Potential contributors should contact the editor
for consideration.
ADVERTISEMENT POLICIES
Advertisements are subject to approval by the editor. The editor reserves the right
to refuse any advertisement. Rates are available upon request from the editor.
BUSINESS CORRESPONDENCE
All communications should be directed to:
Craig Hassapakis, editor and publisher
AMPHIBIAN & REPTILE CONSERVATION
2255 North University Parkway No. 15
Provo, Utah 84604-7506 USA
E-mail: ARC@byu.edu
A Natural History of
Amphibians
ROBERT C. STEBBINS AND NATHAN W. COHEN
This is a book for all readers who want to learn about amphib-
ians. It draws on many years of classroom teaching, laboratory
experience, and field observation by the authors. Robert Stebbins
and Nathan Cohen lead readers on a fascinating odyssey as they
explore some of nature’s most interesting creatures, interspersing their
own observations throughout the book.
A Natural History of Amphibians can serve as
a textbook for students and independent learners,
as an overview of the field for professional scientists
and land managers, and as an engaging introduction
for general readers.
Amphibian populations are being rapidly decimated around the globe, largely due to the encroach-
ment of humans on amphibian habitats and from growing human-caused environmental pollution,
discussed at length in the final chapter. The authors focus our attention on the “natural history” of
amphibians worldwide and emphasize their interactions with their environments over time: where they
live; how they reproduce; how they have been affected by evolutionary processes; what factors will deter-
mine their destinies over time. Through the experienced eyes of the authors, we come to see and under-
stand the place of amphibians in the natural world.
Cloth: $29.95 ISBN 0-691-03281-5
eS,
Lizard Ecology
Historical and Experimental Perspectives
EDITED BY
LAURIE J. VITT AND ERIC R. PIANKA
In this rich collection, leading lizard ecologists demonstrate the
utility of the phylogenetic approach in understanding the evolution of
morphology, physiology, behavior, and life histories. Lizards have been
the subject of reciprocal transplant experiments and of manipulations of
resource availability, habitat structure, population density, and entire sections
of food webs. Such experiments are rapidly rebuilding ecological theories as
they apply to all organisms. As a demonstration of state-of-the-art historical and
experimental research and as a call for philosophical engagement, this volume will
join its predecessors—Lizard Ecology: A Symposium and Lizard Ecology: Studies of a Model
Organism—in directing ecological research for years to come.
“The topics [in this volume] represent an impressive diversity of approaches, rang-
ing from detailed life-history studies of single species and in-depth analyses of lizard commu-
nities to broad-based comparisons of multiple traits across all groups of lizards. . . . [Lizard
Ecology] not only provides a context in which to view these individual studies but opens a
unique window on lizard ecology past, present, and future.”—Science
Cloth: $39.50 ISBN 0-691-03649-7
PRINCETON UNIVERSITY PRESS
AVAILABLE AT FINE BOOKSTORES OR DIRECTLY FROM THE PUBLISHER: 800-777-4726 WORLD WIDE WEB SITE: HTTP: //AAUP.PUPRESS.PRINCETON.EDU/PUPRESS
Amphibian & reptile conservation
: the international journal
devoted to the worldwide
preservation and management of
amphibian and reptilian diversit
HERPETOLOGICAL ECOTOUR AN. MUS. NAT. HIST. LIBRARY
Received on: @8-10-98
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