HARVARD UNIVERSITY
Library of the
Museum of
Comparative Zoology
REAT BASIN NATURALIST MEMOIRS
ber 3 Brigham Young University 1979
MAR 2^1980
The Endangered Species:
A Symposium
GREAT BASIN NATURALIST MEMOIRS
Editor. Stephen L. Wood, Department of Zoology, Brigham Young University, Provo, Utah
84602.
Editorial Board. Kimball T. Harper, Botany; Wilmer W. Tanner, Life Science Museum;
Stanley L. Welsh, Botany; Clayton M. White, Zoology.
Ex Officio Editorial Board Members. A. Lester Allen, Dean, College of Biological and Agricul-
tural Sciences; Ernest L. Olson, Director, Brigham Young University Press, University
Editor.
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iRE AT BASIN NATURALIST MEMORS
U
Brigham Young University
1979
The Endangered Species:
A Symposium
CONTENTS
Introductory remarks. Joseph R. Murphy 1
The epoch of biotic impoverishment. Thomas E. Lovejoy 5
Culture and species endangerment. Roland C. Clement 11
Perspective. John L. Spinks 17
The law and its econimic impact. Donald A. Spencer 25
Endangered animals in Utah and adjacent areas. Douglas Day 35
Endangered and threatened fishes of the West. James E. Deacon 41
Rare aquatic insects, or how valuable are bugs? Richard W. Baumann 65
Endangered and threatened plants of Utah: A case study. Stanley L. Welsh 69
Management programs for plants on federal lands. Duane Atwood 81
Strategies for preservation of rare plants and animals. G. Ledyard Stebbins 87
Strategies for preservation of rare plants. Arthur H. Holmgren 95
Strategies for the preservation of rare animals. Clayton M. White 101
Rare species as examples of plant evolution. G. Ledyard Stebbins 113
The meaning of "rare" and "endangered" in the evolution of western shrubs. Howard C.
Stutz 119
Some reproductive and life history characteristics of rare plants and implications of
management. K. T. Harper 129
The importance of bees and other insect pollinators in maintaining floral species compo-
sition. V. J. Tepedino 139
Endangered species: Costs and benefits. Edwin P. Pister 151
Endangered species on federal lands. Panel: Part I, Introduction. John L. Spinks 159
Panel: Part II, Forest Service philosophy of endangered species management. Jerry P.
Mcllwain 159
Panel: Part III, The Bureau of Land Management's endangered species program. Rich-
ard Vernimen 163
Panel: Part IV, Summary of the endangered plant program in the Bureau of Land Man-
agement. Kenneth G. Walker 165
Index 171
12-79 1.5M 41728
No. 3
Great Basin Naturalist Memoirs
The Endangered Species:
A Symposium
Brigham Young University, Provo, Utah
1979
INTRODUCTORY REMARKS'
Joseph R. Murphy-
As this symposium commences, it is appro-
priate to ask what motivated the conveners
to choose this particular time to address yet
again a now-famiHar subject. More to the
point, what specific aspects of the endan-
gered species problem might be confronted
in the framework of another symposium? At
least three kinds of rationale for such a meet-
ing come to mind.
First, the endangered species program in
the United States, if not throughout the
world, stands at a crossroads in terms of pub-
lic and legislative support. Recent news-
worthy events, wherein an endangered ani-
mal or plant is seemingly pitted against the
agencies of human progress and welfare,
have focused attention on what appear to be
"either-or" alternatives regarding the species
in question as opposed to some real or imag-
ined public good; the Tellicoe Dam incident
exemplifies this kind of dilemma. One result
of this is that industry representatives, and of-
ten government agency personnel, live in
constant fear that some obscure and hitherto
undescribed species of clam or lousewort will
forestall a multimillion dollar development.
Politicians, at both local and national levels,
frequently exacerbate the situation; short on
biology and long on demagoguery, they at-
tempt to undermine the basic concern of the
public for the welfare of endangered or
threatened species. Overzealous con-
servationists may further complicate matters
by adopting inflexible and unrealistic posi-
tions. These polarized opinions leave little
common ground for effective compromise. It
is to be hoped that this symposium can con-
tribute, in at least a small way, to some reso-
lution of these tensions.
A second rationale for the symposium may
be found in the fact that the invited partici-
pants, as well as those who have come to lis-
ten, represent a broad spectrum of agencies
and organizations involved in endangered
species concerns. Among those present are
resource management personnel from numer-
ous state and federal agencies, representa-
tives from public and private utilities and
other industries, university researchers, and
spokesmen for private conservation organiza-
tions. This diversity of viewpoint and expe-
rience should promote a broad examination
of the various issues, and perhaps a greater
tolerance for the views of the "opposition."
Too often, conferences on endangered spe-
cies involve groups with identical or similar
'The symposium convened in the Monte L. Bean Life Science Museum, Brigha
'Department of Zoology, Brigham Young University, Provo, Utah 84602.
Young University, Provo, Utah, 7-8 December 1978.
Great Basin Naturalist Memoirs
No. 3
philosophical positions; this gives partici-
pants an excellent chance to reinforce one
another's views, but little or no opportunity
to gain new insights.
Third, this particular symposium differs
from many held in the past by virtue of its
emphasis upon several taxa of the humbler
and less spectacular creatures, e.g. plants, in-
sects, and other invertebrates. The public im-
age of endangered species is probably embo-
died in or symbolized by such "glamour"
forms as the peregrine falcon or blue whale.
Though strong public support for these spe-
cies is necessary and important, it is mis-
leading and imdesirable to focus all attention
on them alone. Biologists realize that the
condition of the so-called "matrix" species of
plants and invertebrates often supplies a
more accurate indication of the overall
health of an ecosystem than does the plight
of one or a few species of top consumers.
Hence the status of these lower forms be-
comes a matter of priority for all of us, and
not just for the erudite specialist. In a very
real sense, then, this symposium seeks to
break new ground in placing appropriate em-
phasis upon many species which have here-
tofore been neglected.
Obviously, the foregoing are not the only
valid reasons for convening this series of
meetings. There is still a genuine need to ex-
amine the fundamental philosophical prem-
ises vmderlying the management of endan-
gered species. That there yet remain
substantial areas of disagreement concerning
the status of protected organisms was made
manifest in the recent congressional debates
directed toward making significant changes
in the Endangered Species Act itself. In seek-
ing these modifications, congressmen claimed
to be responding to a "grass roots" demand
for relaxation of standards promoted by a
vocal segment of industry representatives and
the public, who contend they have been eco-
nomically disadvantaged by decisions such as
that regarding the snail darter and the Tell-
icoe Dam. The long-term consequences of
any substantial amendments to the present
act will be closely monitored by those on
both sides of controversies involving endan-
gered species.
Broadening the concern for vanishing spe-
cies to a worldwide scope, we find that there
is increasing global concern for the perpetu-
ation of threatened plants and wildlife, as
pressures brought against natural ecosystems
by expanding human populations inexorably
mount. This subject will be treated in great
detail by several of the invited speakers, and
I only wish to point out here that, while
there is much cause for pessimism, there are
a few hopeful signs as well. At the risk of ex-
posing my own biases regarding the relative
worth of threatened wildlife, I will cite a few
positive examples from the realm of avian
conservation, an area in which I have some
first-hand experience.
Certain species of raptors have responded
favorably to the voluntary or enforced de-
cline in the use of persistent pesticides, for-
merly a major source of environmental con-
tamination. Noteworthy in this regard is the
apparent recovery of breeding populations of
the peregrine falcon in the United Kingdom
and parts of northern Europe. A similar pat-
tern of recovery has been detected among
many populations of the osprey in the United
States and elsewhere. It is to be hoped that
this encouraging trend can continue, and that
hard pesticides will be replaced by new-gen-
eration chemicals in those areas of the world
where these problems still exist. In the battle
to save the peregrine, additional successes
have been achieved through the release of
captive-bred birds into areas from which the
species has been extirpated in recent decades.
Results have been sufficiently encouraging to
stimulate the drafting of plans for a similar
effort on behalf of the seriously endangered
California condor; another such program is
contemplated for the Philippine eagle, a vic-
tim of the all-too-familiar story of destmction
of forest habitat to meet the needs of a rapid-
ly growing human population.
The kinds of success obtained with raptors
can certainly be expected in the intensive
management of other species of birds and
threatened wildlife in general. For example,
the International Crane Foundation of Ba-
raboo, Wisconsin, has imderway an ambitious
program of captive breeding and restocking
which has as its objective the perpetuation of
each of the endangered species of that group.
Some additional optimism in respect to the
whooping crane has been engendered by a
scheme to produce more young whoopers for
1979
The Endancered Species: A Symposium
later release into the wild by cross-fostering
of eggs and nestlings, using sandliill cranes as
smrogate parents.
There is another category of species whose
decline may have been arrested by increased
public awareness and more enlightened man-
agement practices of wild populations. Such
would appear to be the case with the golden
eagle in the western United States. While di-
rect and indirect persecution still account for
considerable mortality throughout the West,
there is good evidence that breeding popu-
lations are healthy and stable at the present
time. There are also indications that at least
many of these eagles adjust well or become
habituated to various types of human disturb-
ance, and are not as sensitive or inclined to
abandon nests as many of us had previously
thought. This in turn suggests that not all of
our direst predictions need come true in
every case, although it is obvious that many
endangered species are not as adaptable and
versatile as we would want. Another plus for
the future of the golden eagle is that a long-
awaited federal management plan for this
species appears to be forthcoming. This plan
is intended to offer management guidelines
that will address problems of eagle depreda-
tion as well as deal realistically with the sur-
vival of the species. It is hoped its principal
function will be to keep the golden eagle at a
healthy distance from the endangered species
list; to the extent that it is successful, it could
serve as a model for the management of
many other species.
In closing, I am tempted to deliver a stir-
ring peroration in which I would remind you
of the necessity for coming to grips with the
issues at this critical time in the history of the
conservation movement, of our custodial re-
sponsibility toward the subhuman species
with which we share the planet, and of the
challenge to initiate innovative and effective
solutions. Instead, I will merely quote some
rather straightforward if somewhat trite
words ascribed to the late King George VI,
which were adopted as the motto of the Tim-
bavati Nature Reserve in South Africa:
The wildlife of today is not ours to dispose of as we
please. We have it in trust. We must account for it to
those who come after.
Perhaps it is this intrinsic kind of value
that we should always have before us in our
deliberations on even the least spectacular of
the endangered species.
THE EPOCH OF BIOTIC IMPOVERISHMENT
Thomas E. Lovejoy'
Abstract.— 1978 was the first year in the history of man that legal power to eradicate a species was established. It
is one of a number of signs of rapidly accelerating rates of extinction which may result in reduction of biological
diversity by one-seventh to one-fifth, with a parallel reduction in the planet's capacity to support man and a per-
manent reduction in the potential body of biological knowledge. Species loss of such a degree would warrant desig-
nating the close of the Recent epoch and the opening of a new one of Biotic Impoverishment. A great deal of the
extinctions will occur in the tropical forest areas of the globe but with possible environmental effects extending into
the temperate regions. It will fall to science to help slow the rate of extinction, to decide on which species and
ecosystems to concentrate conservation efforts, and to communicate the importance of biological diversity to govern-
ment and society.
It is an encouraging sign for conservation
that during this first week of December 1978,
both this symposium and the first meeting of
private conservation organizations in Central
America are taking place. It is interesting
that the latter is occurring in Guatemala, a
country which honors the Quetzal, a trogon
of extraordinary beauty, in three ways: as its
national bird, as its monetary imit, and with a
statue in Guatemala City. At the same time
the travel route many of us followed here
takes us through Salt Lake City, where stands
one of the few other statues in honor of a
bird: the gull which rescued the Mormons
from orthopteran plagues, and the specific
identity of which is probably lost forever in
history.
But 1978 will also be remembered as the
year when, for the first time in the history of
civilization, the power to exterminate a spe-
cies other than a pathogen was legally estab-
Ushed. This certainly was not arrived at in
any particularly intelligent manner, and its
full meaning in the history of the biological
degradation of the planet was and is appreci-
ated by few: it is the first indication in the
body of law that we are not going to save, or
try to save, the full array of species in the
biota, and raises the terrifying questions of
which and how many species will be written
off. The new Endangered Species Legislation
takes a step toward answering those questions
by according lower-class status to the faunal
majority represented by the invertebrates, as
opposed to those lucky enough to have spinal
columns.
At first, species will wink out one by one
like city lights as night deepens, but soon
there will be a rushing torrent of extinction.
This year also saw all but the last remnant of
natural forest on Bali cut over, leaving little
natural habitat for the Rothschild's Mynah,
which fortunately does thrive in zoos. As far
as I know, nobody has answered the question
of how many extinctions the Bali forest de-
struction represents. It carries special mean-
ing when we reflect on how much was
learned about how the world works when Al-
fred Russell Wallace crossed the narrow
strait between Bali and Lombok and began to
conceive of the science of zoogeography and,
later, of natural selection. He grasped natural
selection independently of Darwin, yet so ef-
fectively that he propelled Darwin into pub-
lishing the volume which so greatly changed
man's view of his place in nature. It is indeed
likely that some of the recent extinctions on
Bali were of species actually described by
Wallace, a sad tribute to a man who did so
much to advance knowledge.
One little-appreciated aspect of the recent
and forthcoming extinctions is the implica-
tion for the future growth of knowledge
(Lovejoy 1978). An extinct species is one
'World Wildlife Fund, 1601 Connecticut Avenue, NW, Washington, D.C. 20009.
Great Basin Naturalist Memoirs
No. 3
about which we can learn httle, either in
terms of its specific biology or role in nature,
except what can be gleaned from the sad
remnants of information held by museum
specimens— remnants which are nonetheless
valuable and deserving of more appreciation.
Many species will, in fact, disappear without
even a mention of their existence in the
chronicles of science.
While there certainly is some knowledge
to be gained from the response of biological
systems to destructive manipulation, it re-
mains negligible when compared to what can
be learned from living biological systems
over the long time available when species
continue to exist. Indeed one might think of
human knowledge, whether in general or
about biology in particular, as a growing n-
dimensional hypervolume, with a volume Ve
at any point in time. Whether there is a total
potential body of knowledge is probably a
question for lengthy discussion, but for the
moment and for the sake of argument, let us
assume that there is such a definable body,
and that it can be represented by another hy-
pervolume, Vp, which contains, and is many
orders of magnitude larger than, Ve (Fig. 1).
Until relatively recent times, it can be said
that throughout human history society has
striven to enlarge Ve, deriving considerable
benefit from doing so, and essentially driving
Ve to approach Vp. The effects of the rela-
tively small number of man-generated extinc-
tions up to this moment have been minor, but
now, as we enter the epoch of Biotic Impov-
erishment, which prol3ably deserves to be
treated as an epoch distinct from the Recent,
we are allowing Vp to shrink and to ap-
proach Ve. Yet, as living organisms, surely
we must realize that biology is the most im-
portant branch of knowledge for human wel-
fare.
Most of us, even endangered species biolo-
gists, tend to underestimate the extent of the
impoverishment of the biota that may lie be-
fore us. Here in the United States, already a
highly developed nation with a human popu-
lation now predicted to peak at a mere 253
million, it is possible to entertain the notion
that we can have our fauna and flora and the
sybaritic pleasures of the consumer society as
well. There are occasional problems with en-
dangered species and public works projects
in conflict, but it seems possible to at least
dream about having both.
But such is certainly not the case in the
tropical forest countries of the world, which
are, with a couple of exceptions, all lesser-de-
veloped nations eager to ape our ways. The
FAO estimate that tropical forest destruction
currently occurs at 50 acres a minute is truly
terrifying. These forests are a biological
treasury, and the conflicts between tradition-
al development projects and endangered spe-
cies are many orders of magnitude greater
than in temperate regions. I recently had the
Fig. 1. The relationship of expected (Ve) and poten-
tial (Vp) knowledge.
1979
The Endangered Species: A Symi
difficult task of producing for a presidential
study an estimate of extinctions that will take
place between now and the end of the cen-
tury. Attempting to be conservative wher-
ever possible, I still came up with a reduction
of global diversity between one-seventh and
one-fifth, principally because of what will
happen to tropical forests. No doubt some of
my colleagues think I am mad, but I would
challenge them to produce a better estimate.
If my estimate turns out to be too high it will
either be because society has made consid-
erable changes in its ways, or because it will
take a bit longer to reach the same reduction
in diversity. The U.S. example to the inter-
national community will be very important.
The problem of biotic impoverishment
then is considerably greater than can be ap-
preciated from an overview of endangered
species here in the United States. Our Ameri-
can collection of endangered flora and fauna
really represents but part of the forward con-
tingent of a great rush to extinction. This all
raises many questions for science and society,
including the most terrible one of all, one
which few are brave or foolhardy enough to
ask: namely, can we continue to treat the val-
ue of a human life as a constant whatever the
number of people may be? I am enormously
uncomfortable even asking it, let alone trying
to answer it, yet, even if ignored, it will in
part be answered by the degraded capacity
of the planet to support man if biotic impov-
erishment proceeds apace.
It is more comfortable to try and answer
the scientific and technical questions raised
by biotic impoverishment and it is these that
this symposium is largely addressing. Cer-
tainly science has a critical role to play in ef-
forts to reverse the tide, and it needs to be
recognized that such science is as in-
tellectually respectable and as useful to so-
ciety as, for example, laboratory work on
DNA hybridization.
In many cases it is a race with time. World
Wildlife Fund is launching a major research
program on the problem of the minimum
critical size of ecosystems, or why and how
ecosystems set aside in the midst of land-
scapes converted to man's purposes shed or
lose species. How much more desirable it
would be to have the results in hand now, be-
cause most opportunities to set aside a repre-
sentative series of the planet's ecosystems
will occur in the next two decades. Yet we
have to make do with the situation.
It is important that universities and
agencies recognize that conservation is an
appropriate activity to which biologists
should devote some of their time. Indeed it is
probably correct to say that today a biologist
can make as much of a contribution to sci-
ence by helping to save species and ecosys-
tems, and, therefore, future opportunities to
study biology, as by more traditional scholar-
ly pursuits.
To science should fall the terrible questions
about which species and ecosystems to save
and which not to save. Which of us would be
comfortable about saying one species is more
important than another? Yet it is clear that
all will not be saved, and that it is preferable
for scientists to address those questions rather
than deferring to the less knowledgeable.
Certainly we would be wise to try and avoid
having to make such decisions in imthought-
ful haste— but maybe the awful nature of the
questions will drive us that much harder to
keep the number of such decisions small.
There is another critical role for science,
namely articulating the true meaning of biot-
ic impoverishment for society. Too often
both scientists and society in general focus on
the individual species rather than on the rec-
ognition that the impoverishment of the
biota represents a reduction of the planet's
capacity to svipport man. Just on the basis of
destroyed coastal wetlands it can be said that
the fimdamental capacity of the planet to
support man is less today than a century ago.
This erosion of our biological wealth has
been masked by the constantly refined abili-
ties of technology. Technology has fostered
the illusion that we can get more and more
from less and less, a dream that soon will be
shattered; the cracks are already appearing.
Scientists must articulate the true meaning
of endangered species as indicators of
stressed ecosystems and as yet another sign of
erosion of the basic quality of life. When the
Devil's Hole pupfish was being endangered
by a lowered water table, the real question
was about the rights to reduce that natural
resource base. It is not always easy to deduce
the complete meaning for society of any par-
ticular endangered species, but it will always
Great Basin Naturalist Memoirs
No. 3
be generally true that it will reflect deterio-
ration of a biological system. When arguing
the case, we cannot always expect the exploi-
ters to play fair; poor human behavior with
respect to questions of dwindling resources is
all too familiar.
Our society is generally ignorant of the im-
portance of biological resources and hence of
endangered species. One Supreme Court
judge harbored the view during the case of
the Tellico Dam and the snail darter that the
Endangered Species Act meant endangered
species might be found behind already exist-
ing dams and it would be necessary to run
around letting the water out. And a distin-
guished senator was concerned most of his
state might be declared critical habitat for
the grizzly bear; perhaps he was worried
about their votes? It is really up to all of us
to employ all our energies and intellects to
correcting this dangerous state of ignorance.
We should clearly make the case, for ex-
ample, that dwindling biological resources
are making a major contribution to problems
of economic inflation.
Society worldwide has been rocked in re-
cent weeks as the story of the charnal mille-
feuille of Jonestown emerged layer by layer
in the media. How many of us recognize that
we are well into the beginnings of a biologi-
cal Jonestown? As in Jonestown, there will be
human survivors once the epoch of biotic im-
poverishment passes. But their existence will
be forever an impoverished and degraded
one. The question of how degraded and how
impoverished lies before science and society.
Acknowledgments
I acknowledge with gratitude the help of
Richard O. Bierregaard, Helen Hays, Eleanor
Stickney, and Clayton White.
Questions to Mr. Lovejoy
Q. It would be interesting to have a figure on that one-
sixth of the species lost you have computed.
A. You mean in terms of numbers of species? If the to-
tal biota for the planet is estimated at lying some-
where between .3 million and 10 million species,
which gives you some idea of how ignorant we are
about the biology of the planet, and if one takes the
more conservative figure of 3 million, the loss is on
the order of 500,000 to 600,000. The actual home-
work I went through to arrive at all of that is going
to be published in the Global 2000 Study, which was
requested by President Carter more than a year ago
in his environmental address of 1977 and will
emerge sometime in the coming September. That
particular section will certainly stand out because a
lot of the reviewers of that study who are not biolo-
gists and not particularly aware of whats happening
in the tropical regions found it very difficult to be-
lieve and so it may be the only portion of that study
with a name actually attached as an author.
Q. Would you explain your statement about fauna
without vertebrae having less of a status in nature
than vertebrates?
A. Well it's true. When it comes to invertebrates the
secretary of interior has the ability to exempt certain
public works projects from the Endangered Species
Act without taking it through the long process to ul-
timately what some of us call the "Extinction
Squad" setup in the new legislation. It really doesn't
say flat out that the secretary of interior can allow a
nonvertebrate to go extinct, but it says that he need
not recognize discrete populations. Maybe some of
the Department of the Interior people can explain it
a little more fully than I, but there was a clear dis-
tinction drawn between the two.
Q. What are the strategies to address the problem of
the destruction of tropical rainforests?
A. As currently treated, tropical rain forests are gener-
ally regarded as one-time use resources to be cleared
and replaced by something else. That something else
is often not economically viable in the long term. It's
really a matter of proving that there are in fact con-
siderably more gains from protecting as much of the
tropical rain forests as we can than from converting
them to short-time use.
Q. The recent modification of the Endangered Species
Act was supported around the country. Some of the
major conservation groups supported that modifica-
tion. Now I understand that the World Wildlife
Fund also supported the modification and I'd like to
know if that's correct or what your position is.
A. During the period when the new endangered species
legislation was being considered in recent months,
some conservation organizations supported the mod-
ification. Was World Wildlife Fund one of these? It
was not. The problem was that the biological reality
and the political reality simply did not coincide. It is
very clear, particularly from the Senate vote, which
I think was something on the order of 94 in favor of
modification, that there was little sympathy within
the Congress for maintaining the Endangered Spe-
cies Act as originally laid out. We, in fact. World
Wildlife Fund and some other organizations, worked
quietly behind the scenes to try and get as strong
successor legislation as we could possibly get, but we
simply were not about to go on record and condone
anything that might lead to extinction. I think this
was well understood by the staff of the Senate sub-
committee in question.
Q. I have two comments to what you had to say today.
First of all, forgive me if I have trouble thinking
about tropical rain forests today. I deal in the area of
management, which is where most of the problems
1979
The Endangered Species: A Symposium
are being solved, at least in this country. If we set
ourselves as biologists aside and say it's iis against
them, then two things may happen. First, we may
become like a federal agency that I know of that
says we have this pristine untouched piece of land
and we're going to keep it pristine and untouched
by allowing 10,000 people in and then it's going to
stay somehow miraculously pristine and untouched.
Second, which is more likely, the "other side" will
win because they have more influence and more
people. I think that we need to concentrate more
heavily on the interface between the scientific com-
munity and the people and, in a democratic country,
I think that is more appropriate.
\. I don't have any difficulty with that at all. In fact, I
think in a sense that's what I was trying to say by
saying we have to articulate what it all really means.
But it's all in the best interests of all of us to be wor-
rying about what is essentially the proper biological
management, not only of the United States, but of
the entire planet and that naturally falls into two
distinct parts. One is making sure that you protect
all the pieces— all the biological pieces— with a series
of representative ecosystems properly designed so
that thev can be managed to protect their integrity,
and I might add that it's getting harder and harder
to protect an ecosystem anywhere without manage-
ment of some sort. The other part is making the rest
of the face of the planet biologically productive in a
sustainable fashion and in a fashion that will in no
way threaten the sort of species bank you've set
aside. For example, we at World Wildlife Fund are
taking a long shot at trying to talk to the last of the
American billionaires, Daniel Ludwig, who has an
operation in the Amazon about the size of Con-
necticut where he has cleared forest and is growing
trees for pulp and rice. If I actually do get a chance
to have a reasonable discussion with him, my point
to him is not going to be that he shouldn't be doing
what he's doing, because it is probably more in-
telligent than most of what's being done in the Ama-
zon, but rather that the long-term security of his
own operation depends on protecting the ecological
integrity of the Amazon as a whole.
Q. You alluded to a correlation between biotic impov-
erishment and economic inflation. Would you care
to elaborate on that subject?
A. Probably, if one searched around, examples could be
found. For instance, the price of fish is increasing as
the supply dwindles and the demand stays high. I
only began to think about this ten days ago, but cer-
tainly this process has to be occurring here and
there. The problem is to sort it out from the incred-
ible maze of vectors that comprise economics and
really demonstrate what's happening— and that's
part of the problem.
Q. Mammoth is now about $5 a poiuid. The summer
Chinook is in very great danger; we might have to
drain Grand Coulee Dam to save it. In one of your
other statements, you said that we have to protect
habitats that have been damaged or destroyed and
renew them. No summer Chinook were caught this
year, except for about 1,.500 by the Indians. At one
time there were billions of pounds of Chinook. Crab
is now about $5 a poimd.
A. I'm really delighted to have that example. Well, in
fact, if you probably start thinking about it, the
price of lobsters and other marine products has cer-
tainly risen a great deal in recent years and that may
be one of the more clear-cut situations. The world
fisheries peaked about 1970. They made a slight re-
covery this year, I believe, but all along there has
been an increasing catch effort, an increasing
amount of actual fossil fuel energy going into the
pursuit of these fish, and certainly the demand for
them has been increasing, too.
Q. Are there any calculations on how much of a de-
crease we are going to get in atmospheric oxygen if
these tropical forests are destroyed?
A. The problem of tropical forests with respect to oxy-
gen turns out to be no real issue. They, in fact, con-
sume about as much oxygen as they produce. The
real problem in terms of biogeochemical cycles may
well lie in the carbon which is stored in tropical rain
forests. Tropical rain forests represent the largest
terrestrially stored pool of carbon, and it was esti-
mated as long ago as 1954 by Evelyn Hutchinson
that the increase in carbon dioxide in the atmo-
sphere probably came equally from the burning of
fossil fuels and the destruction of forests. The whole
problem of global circulation of carbon and where
the sinks are certainly isn't a clear matter, but we do
know that CO2 is increasing and that if we destroy
two-thirds of the tropical forest (which is the esti-
mate by 2000) we will release an enormous amount
of carbon into the atmosphere. The question is how
rapidly the natural sinks can absorb it and bring it
back to normal level. If there is a pulse of carbon in
the atmosphere, then we may get into problems of
climatic change, change in rainfall regimens, and
temperatures in the temperate zones. The tropical
rain forests really aren't all that far away.
Q. It seems like we're expanding on the strategy of the
common ideal. How can we deal with individual
countries that are bent on destroying a habitat
which may affect the rest of us?
A. If I had the answer to that I might be head of the
U.N. There is no easy answer to it at the moment.
Many, such as Roland Clement or I, work on a coun-
try-by-country basis. I endeavor to persuade the Bra-
zilians, for example, that proper management of the
Amazon is in their best interests as well as that of
the rest of the world in terms of, say, carbon. On the
other hand, there is increasing recognition within
governments, even in Latin America, which people
were despairing about for so long, that there are ma-
jor problems. There is now an Amazonian pact
among all the Amazonian nations which, at least in
rhetoric, talks about proper management of the
Amazon. There are some interesting questions in-
volved in that. Take the example of the contribu-
tions of biological species to modern medicine, espe-
cially the importance of a primate species. Take a
cure that comes from one of those species and it is
verv rapidly distributed around the world as an in-
ternational resource, so we may be approaching the
time when some biological resources, at least, are
10
Great Basin Naturalist Memoirs
No. 3
recognized by the international community as being
international resources, and pressure will therefore
fall on some of the less well-behaved countries to
clean up their act.
Q. Are people as much a problem as governments?
A. Oh, it certainly is both. Roland Clement is going to
address problems of people.
Q. Is corruption ever a problem?
\. Well, corruption is a problem. I would guess that at
least in Latin .\merica, which is my major "beat,"
ignorance is a greater problem than corruption.
Q. Do you think there really is a middle ground be-
tween biotic deprivation and the goals of preserva-
tion of the biota?
A. I think there has to be. It's just a matter of how
much we want to let it hurt before we really re-
spond to it. 1 think it's as simple as that.
Q. What about the review board set up under the new
endangered species legislation to make decisions on
seemingly irresolvable conflict between endangered
species and public works projects? How busy will
they be, and how eager will they be to exercise that
power?
A. We're really talking about a play of attitude and
how that will affect the whole process. The Endan-
gered Species Act up to the point where it was re-
vised certainly involved thousands of cases of seem-
ing conflict that were all resolvable at staff-level
discussion with the Department of the Interior. The
single exception was the Tellico Dam and snail dart-
er, and that really became an exception only be-
cause the TVA was completely recalcitrant and re-
fused to acknowledge that it was subject to the law.
If one reflects on those figures, it might say that only
rarely will the Extinction Squad have a nasty deci-
sion to make. If, on the other hand, agencies feel
that the creation of the Extinction Squad has weak-
ened the power of the Department of the Interior,
they may feel less need to be pliable in discussion
and they may get very busy up on top. I don't think
they're going to be happy to do this at all. My own
particular .solution to the whole thing is to erect a
large black marble slab on the mall in Washington
so that we can engrave on it the name of the species
exterminated and the names of the members of the
squad at that time.
Literature Cited
LovEjoY. 1978. The Curious Animal, Symposium on
"Impact of Federal Wildlife Regulations on the
Svstematics Ecology Community," sponsored by
the Society of Systematic Zoology and the So-
ciety of Plant Taxonomists, 14 Febniary 1978.
CULTURE AND SPECIES ENDANGERMENT
Roland C. Clement'
.\bstract.— Species endangerment has so far been addressed mostly by biologists. It is now important to involve
social scientists, inasmuch as the problems are man caused. The history of our attitudes, our uses of the land, and the
reasons wherefore are problems for everyone.
The evidence suggests that the causes of endangerment may be grouped under (1) direct and indirect exploitation
of resources, and (2) population displacement by modern agriculture, with consequent migration to the city or to the
forest frontier, where accelerated forest destruction is the result. The displaced people are part of the marginalized
two-thirds of the human race and will destroy what is left of nature in order to survive unless we help them become
self-sufficient.
Such a refocusing of Western civilization, which has so far been parasitic on nature and a marginalized humanity,
will require a new world view by the dominant one-third of us, perhaps based on Whiteheadean philosophy, wherein
we accept a participatory role in a complex of processes that evolve from one another.
The focus of this paper is an attempt to
broaden perspectives on contemporary spe-
cies endangerment. The biology that is cur-
rently being elucidated by a spate of dis-
cussions of this problem is fascinating and
helpful, but not enough in itself.
Let us first summarize very quickly the
two principal causes of man-caused endan-
germent under two categories of pressure: ex-
ploitation and competitive exclusion. These
are well known, but we tend to generalize
them too much. For example, there is abun-
dant and growing objection to direct exploita-
tion, such as whaling, sealing, even hunting,
and more recently against economic devel-
opment which destroys key habitats. But we
still neglect the impact of the killer-buyer
relationship in the exploitation of wild spe-
cies, partly because it is diffuse and largely il-
legal, and therefore difficult to quantify. It is
also a more recent phenomenon. The new
traffic in animals and their parts We need to
confront and regulate is a by-product of the
jet age and the mass-consumption society. It
is a result of uneducated affluence.
There is cause to believe that the United
States alone generates a $10 million annual
traffic in live birds; that two to four times the
number of individuals delivered perish en
route; and that "products" made from wild
animals involve sales which are several times
$10 million. The Justice Department is now
attempting to assess this traffic more accu-
rately, almost for the first time.
There is also a need to study the implica-
tions of the sheer weight of human numbers
on management policy. Numbers probably
now mitigate against the rational manage-
ment of our wildlife resources. What we re-
cently considered "moderate use" now adds
up to excessive demand and exploitation.
It is the same with competitive exclusion.
We stress human population growth and tend
to point the finger at the poor who still favor
large famlies. But we have neglected the so-
cially disruptive displacement of people by
agricultural "modernization." Thirty years
ago, the growth of capital-intensive and tech-
nology-intensive mechanization in U.S. agri-
culture sent 10 million blacks to the cities,
and today the Green Revolution or some less
spectacular form of agricultural modern-
ization is doing the same thing to world peas-
antry. The only way to call this modern-
ization progressive is to overlook the social
and ecological disruptions for which it con-
tinues to be responsible.
The displaced people are flooding the
cities and rapidly destroying their viability
because the cities are incapable of assimilat-
ing such numbers. Or the people become
modern-day colonists along the frontiers of
R2, Norwalk. Connecticut. 068.50.
11
12
Great Basin Naturalist Memoirs
No. 3
the remaining forest, destroying it at a ca-
tastrophic rate.
Having been slow to understand the social
dynamics of labor displacement by machines
and gasoline, we have misidentified the im-
pact of marginalized human groups. We
blame what seem to us insensitive attitudes
toward forest destruction by these people, or
we point to over-population as a basic cause.
We must learn to recognize that today's for-
est destruction is a by-product of our own
economic demands. It was to facilitate min-
ing activities by foreign capital that Brazil's
trans-Amazon highway network was built;
and it is to satisfy America's hamburger cul-
ture that cattlemen have displaced corn
farmers from nearly half of Central America.
Another neglected element of competitive
exclusion which needs to be seen as a sense-
less ecological pressure is that of the mecha-
nized mobility of this generation. This is ex-
hilarating, but it allows one species to disrupt
the existence of all other species as never be-
fore. Its unfavorable effects are another result
of uneducated affluence. This effect may, of
course, be constrained within a decade or so
by the energy shortage.
At a La Jolla seminar on endangered spe-
cies, Lovejoy (1978) outlined the scientific
needs of late twentieth-century conservation.
He called for species research to identify
minimum habitat requirements, for island
biogeography-type studies to identify species
maintenance needs as against individual
needs, and for ecosystem research. Lovejoy
also called for a new conservation anthropol-
ogy focused on the study of human attitudes
and values, but leaving out the needs of the
Third World, important as he knew these to
be. He asked that this new field of study in-
quire into the biology of our own species, so
that we may learn what has led us to take
such an adversary stance toward the environ-
ment.
As already suggested, much as I would val-
ue more biological knowledge of ourselves
and other species, this will not suffice to re-
duce the rate of extermination which is now
underway. We need a conservation anthro-
pology but it must be a much broader inquiry
than attitudinal res-earch; it must investigate
our world view, which is the way people
"characteristically look outward upon the
universe." This calls for an investigation of
our culture, because culture is the sum of our
ideas about ourselves, our environment, and
the social institutions we have devised to get
things done.
Anthropologists (Hall 1977) tell us that we
cannot understand our own culture by self-
examination or introspection, but only by
comparing the approaches of other cultures.
Other cultures furnish us a necessary point of
reference. Therefore, rather than the psy-
chologist, it is the historian, studying older
cultures, and the anthropologist and the ge-
ographer, systematically studying existing
cultures, who are our best guides to under-
standing ourselves and our neighbors. They
can help us unravel the social psychoses that
cause us to undermine our own existence
through internecine struggles and the impov-
erishment of the biosphere. How ironic that
we have heretofore been so heedless of the
survival of other cultures. We need one an-
other, if only for dead-reckoning purposes!
Will it turn out to be the same with those
other nations, the wild species?
Our own culture, a variant of Western civ-
ilization, is now old enough to be viewed
somewhat objectively by mature historians.
Ordinary citizens cannot yet do this because
they have internalized culture and are not
yet scholarly enough. But they can be taught.
Two of the most intriguing historians of our
day who may serve as models and teachers
are Fernand Braudel (1978), the Frenchman
who is the authority on the growth of civ-
ilization in the Mediterranean basin, and Im-
manuel Wallerstein (1974), who is engaged
on a three-volume study of the modern world
system which is our economic system.
The particular importance of Wallerstein
for our theme is that he has disaggregated the
modern world system. We are all familiar
with the fact that the nations of the world
normally include one or a few dominants, but
Wallerstein is convincing in demonstrating
that this is an interdependent system, where-
in a small group of core nations system-
atically exploit a larger group of peripheral
nations. The danger is that the core nations
build up such a high life-style by exploiting
the peripheral nations that they become de-
pendent on this unbalanced system of ex-
changes. It is noteworthy that it is this sub-
1979
The Endangered Species: A Symposium
13
sidized well-being that allows the core
nations to enjoy the internal political free-
dom they boast about and mistakenly see as
the cause of their well-being instead of a by-
product. The peripheral nations, conversely,
must impose some form of coerced labor to
extract the raw materials for export, while
consumption at home is constrained. Eco-
nomic development for the core nations is
maintained by underdevelopment of the per-
ipheral nations. This is a large part of the
Third World problem.
History also teaches that dominance in this
system is a shifting phenomenon. Spain was
the first leader, but for less than a century.
The British enjoyed dominance for a very
long time, but lost it to the United States dur-
ing World War II. There are many signs that
the dominance of the United States may not
be longer lived than Spain's unless we be-
come more appreciative of the relationships
involved and modify the system to remove its
worst inequities.
In any event, because the most worrisome
threats of extinction are focused in the
world's tropical regions, and these areas are
all peripheral nations in the world economic
system, it is obvious that we must look to the
exploitive life-styles of the core nations if we
are to introduce a more rational balance be-
tween numbers of people, their material de-
mands, and the carrying capacity of the re-
gions involved. It is the mass-consumption
society that has the most "give" in it. A first
task in this monumental transition may be to
analyze and display those ecologically sense-
less agencies in our culture that have provid-
ed the compulsive drive to exploit the planet
as though it were a midden heap. People
must be shown that their culture has become
counterproductive before they can be ex-
pected to make the necessary changes.
In addressing the particular problems of
the Neotropics, we will benefit from Brazi-
lian anthropologist Darcy Ribeiro's (1971)
analysis of our Western Hemisphere system.
The cultural complexity of the hemisphere is
highlighted by Ribeiro's need to recognize
three categories of people. First are the Wit-
ness People, the descendants of earlier native
civilizations that were crushed by the Span-
ish invaders. These are the Aztecs of Mexico,
the Maya of Guatemala and environs, and the
Incas of Peru and Bolivia. These people com-
prise a large but marginalized component of
the modern-day populations of their new
countries. A second group is that of the New
People, where the European colonists
amalgamated lesser tribes, including im-
ported African slaves, and thereby created a
new mixed human stock. Such an amalgam is
characteristic of Brazil in particular, and of
Venezuela, Colombia, Chile, the Antilles,
southern Central America, and the southern
United States.
Finally, there are two groups of Trans-
planted People, New World Europeans so-to-
speak, who simply pushed native peoples
aside on first conquering their areas. These
are the Anglo-Americans of Canada and the
United States; and the River Plate people of
Argentina, Uruguay, and Paraguay. Notice
that these two groups occupy the temperate
zones of our hemisphere. We are still, in
many respects, a bunch of tribes, but we are
tied together by an economic system whose
exploitiveness has become unbearable for
two-thirds of the world population newly en-
lightened about their true status by American
movies, radio, and television.
Let me be more specific by calling atten-
tion to recent studies of what is happening in
Central America. In what is unfortunately a
somewhat obscure publication for many of
us, Berkeley geographer James J. Parsons
(1976) reported that the expansion of arti-
ficial pastures in Central America at the ex-
pense of both cropland and natural forest is a
regional phenomenon with drastic, over-
looked consequences. In most of this region
in the 15 years prior to Parson's report, the
area in planted pasture (mostly in African
grasses) and the number of beef cattle had
nearly doubled. But the per-capita con-
sumption of beef in those same countries had
actually declined, because most of this pro-
duction was for export. This expansion of
cattle raising was done at the expense of the
Indians and mestizos who formerly raised
corn for themselves on these marginal lands.
Dispossessed by the cattlemen, these people
have either migrated to the cities or have
gone to the forest frontier to engage in shift-
ing cultivation by cutting the forest. Within a
few years, of course, the colonists find it nec-
essary to abandon their plots and to cut new
14
Great Basin Naturalist Memoirs
No. 3
forest, mostly because weeds become more
expensive to fight than cutting forest. The
cattleman then moves in behind them, rents
the abandoned acreage for a pittance, and
plants it to grass. The peasants have not only
been dispossessed, but they have become a
free labor supply for cutting the forest.
Although lauded as progressive and mod-
ernizing by local and national governments
and bv international agencies like AID and
FAO, it is this system of land use which has
become the chief cause of tropical forest de-
struction in the neotropics. It is, of course,
also impoverishing already marginalized hu-
man populations. We have all known for
years that the pressure of so-called land
squatters, with machetes and fire, were a
serious threat to any forest preserve in Latin
America, but Parsons and a few young an-
thropologists were the first to show that this
destruction is the end product of a wide-
spread economic system anchored in the
hamburger culture of the United States. The
U.S. link has not yet been spelled out in de-
tail, but it seems obvious. The dire effects of
this extensive land use shift of the last 20
years or so on the Indians of Chiapas are now
being documented by Robert Wasserstrom
(1977, 1978), James D. Nations (1979), and
their colleagvies who worked, until recently,
as the Centro de Investigaciones Ecologicas
del Sureste (CIES) in San Cristobal de Las
Casas.
It is time to suggest that the environmental
awareness that has come to so many in this
decade of the seventies is akin to a religious
revelation. Having become aware that our
economic system is parasitic on both nature
and people, we are now challenged to redes-
ign our world view in line with a more con-
sistent vision of the joint realities of our lives:
the environmental, which is the substrate of
our existence; and the social, which is a mea-
sure of our humanity in making intelligent,
perhaps even enobling use of our opportu-
nities. This calls for a reassessment of our sci-
ence and technology, our values, and even
our unstated, unexamined theological as-
sumptions, since willy-nilly, we have some
vague concept of the destiny of mankind.
Obviously it must be a joint venture, and it
will take time, but each of us can help by
engaging in some fraction of the task and by
involving others.
We are likely to discover in that process of
review that the principal assiunptions of our
Western civilization— homocentricity, ration-
ality, technocracy, and progress— have be-
come an embarrassing myth. We have ido-
lized our own creations instead of simply
appreciating them as events in our history as
developing organisms. Having demytholo-
gized nature and his origins, modern man
himself now stands in need of demythologiz-
ing!
Ecology has taught us that we are involved
in systems within systems, and that we both
impact and are impacted by these systems.
But scientific reductionism, useful though it
may be as methodology, has become a dan-
gerous, unwitting philosophy. It seems likely
that a great deal of that sense of relationship
to the environment which we lack, but which
the ancients had, is due to the specialism and
incrementalism encouraged by reductionism.
This has, of course, also affected our educa-
tional approaches and made for a hasty em-
phasis on specialization for the sake of pre-
paring practitioners. Education should
involve helping people see whole systems be-
fore training them to analyze and manipulate
the elements of these systems.
However, to invent a new outlook is not to
destroy the old, but to give it a new form, a
new emphasis, a new reach. Jay Forrester
(1978), who prepared the way for the Limits
to Growth debate, has now suggested that we
have perhaps already been through the tech-
nological age. This does not mean that we
are through with technology, but that the age
which is dawning will not be awed by tech-
nology and will use it in the service of all
men instead of as an end in itself and for a
relative few.
It seems obvious that, if we are to save the
million or so species we fear may be lost
along with the destruction of the tropical for-
ests of the world, we must open our system of
inventive production to that two-thirds of the
human race which was marginalized during
the mad rush for domination. The margina-
lized people will otherwise be forced to chew
up the forest in a frantic effort to survive.
But the earth cannot support its present
overload of humans at a standard of living we
1979
The Endangered Species: A Symposium
15
would like. The first step must therefore be
to eliminate waste in order to make our re-
sources satisfy more people; then to tailor
our demands to more modest proportions;
and finally to adjust our numbers to a new
sustained yield economy instead of the pres-
ent liquidation of resources that passes for
production. Done in stepwise fashion, we will
be pleased to see that efficiency and frugality
do not hurt.
We can draw inspiration from the process
philosophy of A. N. Whitehead (1933) of a
half century ago, and from the new interest
in the implications of historical con-
sciousness. Whitehead's cosmology is summa-
rized in the thesis that "the ultimate and fim-
damental reality of the universe is a multiple
and never ending complex of processes devel-
oping out of one another." This is both a sci-
entific and a metaphysical statement of fact.
To think in process terms is to acknowledge
our dependence on the systems that produce
us and our responsibility to contribute, in-
sofar as we can, to the advancement of these
processes instead of destroying them for tem-
porary self-satisfaction. The way of historical
consiousness (Stevenson 1969) is a restate-
ment of the same concept: that when the
reality of existence, and we ourselves, are un-
derstood as historical, we become aware of a
responsibility to and for history. In both cases
the appeal is not to morality as injunction,
but to participation in a process.
The theologian Paul Tillich once said that
the salvation of man and nature are one and
the same task. More recently the anthropo-
logist Edward T. Hall said that the popu-
lation-environment crisis and the crisis of
relationship to self must be solved together.
It seems to me we have enough testimony to
get started on the reconstruction of our cul-
ture.
Postscript
A frequent response to the approach taken
in this paper is that it is too optimistic, as
though I expected things to right themselves
as soon as awareness is more widespread. It is
also objected that "education" takes too long.
But education is not restricted to that long
sequence of school attendance we currently
impose on our young. It may also affect those
in control of our social systems, and, through
them, all those in between. A culture does
not change until all the people in it also
change. There is no telling how long this will
take, but a crisis or unusual leadership may
make it happen rather suddenly.
The destructive portend of current prac-
tices has caused British astronomer Fred
Hoyle (1977) to suggest that the salvation of
the human race may depend on an early col-
lapse of our economic system. He sees two
likely options for a high-technology society
like ours: (1) if an essentially unlimited source
of energy were perfected before the human
race agrees to limit its population and subsist
by less destructive life styles, a collapse lead-
ing even to extinction is likely. If, on the oth-
er hand (2) an early economic collapse causes
us to come to terms with ourselves, and we
limit population everywhere, the consequent
rebirth of invention, if it then provides ample
energy supplies, may allow the human race
to rise to new heights that are hardly imagi-
nable at present.
Questions to Dr. Clement
Q. How can we show people that their culture has be-
come counterproductive and needs to be changed?
A. Let me first make the point that cultures don't
change until almost everybody changes. It is an
educational process, and scientists will need to help
by pointing out the implications of what ecology is
teaching us; that way we will revamp our education-
al system. If the people don't understand that they
partake of larger systems, they will continue the
short-sighted exploitation which has characterized
our civilization. We always begin by accepting the
cultural systems we are born into. And it was a
great, exciting, and in one sense enlarging expe-
rience to be caught up in this wave of exploitation.
But now the very system of exploitation is in ques-
tion, so we must help people understand that we are
not proposing a Marxist revolution but a revamping
of our system before it breaks down. There are abun-
dant signs that the breakdown is already underway.
Q. Will we succeed in revamping our civilization to
prevent the extinctions you and Lovejoy are so con-
cerned about?
A. Let me take a different tack. Education usually takes
a generation, but it may come quickly if a crisis oc-
curs and our leaders can point out new directions;
the people may then ttim around almost overnight.
One reason I'm optimistic about the future is that
our system is so close to its end that we will not
achieve the growth projections in which the business
world believes. We are already so deadended in so
many areas that if we don't wake up to our problems
16
Great Basin Naturalist Memoirs
No. 3
in a decade or so, we'll be squeezed down to size. It
would take another lecture to develop this point,
but the petroleum problem is a good clue. We have
a naive faith that all we need do is invest more in
production at home to get more energy. Of course
we can do this, but only at increasing cost. Every
million feet of new oil well costs more than it did
yesterday, both in dollars sunk and in diminished re-
turn. And the more we pump, the quicker we will
run out. This is what needs to be made obvious to
the people. We currently buy the surpluses of the
Middle East because these are the cheapest ones.
Even American agriculture will have to be turned
around because of the energy crisis.
Q. You talked about marginalized people in Latin
America, but are there not marginalized people in
the U.S. who will get caught in this crossfire?
A. There are many other marginalized people— the
blacks, the hispanics, and the Amerindians at home,
and the people of Asia and Africa. Reassimilating
these people into the world economic system will
not be done overnight, but if we at least accept this
as a challenge and work at it, we will soften the im-
pacts.
Q. What kind of progress is Canada making toward be-
coming a conservationist society?
A. I'm afraid they are not making much. The Cana-
dians are making all the same mistakes in exploiting
their energy resources that we have made. This is
not surprising because Canadians are a marginalized
people too; they are exploited by the United States.
Q. You spoke of the need to educate the public to the
fact that our system has serious problems; you also
said that we must convince people that they will
need to get by on a smaller piece of the resource pie.
How can you really expect this to work? Won't some
economic penalty be necessary? People usually
change when they see a personal payoff of some
kind.
A. Well, yes, but you are opening up another area
which cannot be addressed with a simple answer.
There are no simple answers. In a democratic so-
ciety we must seek to persuade rather than impose.
But if time runs out we face a dilemma. Within a
decade or so we will probably truly be in an energy
crunch, and we will then learn that volimtary ap-
proaches to the conservation of energy are not
enough; they put the good guys at the mercy of the
cheaters. When that time comes, it is hoped some-
one will have the persuasive skills to put this over. If
we don't accept this, we'll have to fight over it; and
if we do that we're in real trouble.
Q. Isn't that happening now? This audience is aware of
many of these problems, but we also have large cor-
porations with lots of money advertising in national
magazines and on television, saying: "Don't worry.
Buy your gas dryers and gas stoves. We'll have gas
for as long as we need it."
A. Exactly. We already have a conflict of approaches. \
large section of the economic community actually
still believes that resource "production" is simply a
matter of investment. Look at the ads nm by Mobil
Corporation, "The Capitalist Revolution." They say,
just get the government off our backs and everything
will be all right. They seem ignorant of the fact that
they are liquidating the resources. Of course, if you
don't care about the future, that's another matter.
Q. Isn't it correct that increasing the price of, say, pe-
troleum may not solve problems of exhaustion and
inflation?
A. Yes, but price is important. If we can make people
pay the full social costs of what they wish to do,
they must then decide what they most wish to do in-
stead of greedily trying to do everything, or con-
suming everything. If we have artificial price struc-
tures, the public is misled. Industry agrees with this
view. The question is who will get the price in-
crease? It should be a tax that we can use for con-
stnictive uses— in the case of energy, to rebuild a
mass transit system where the people are.
Let me now add that I'm delighted at the re-
sponse you provided because a large part of the an-
swer to this big problem is people like you tackling
questions energetically and in an open fashion. We
must then try to involve more people in our tenta-
tive conclusions. If we don't, our troubles won't go
away.
Literature Cited
Braudel, F. 1978. Afterthoughts on material civilization
and capitalism. Johns Hopkins.
Burger, J. M. 1965. Experience and conceptual activity.
MIT Press.
Forrester, J. W. 1978. Changing economic patterns.
Technology Review, Aug. /Sept.
Hall, E. T. 1977. Beyond culture. Anchor Books.
Hoyle, F. 1977. Everyman's universe. In: Ten faces of
the universe. W. H. Freeman and Co., San Fran-
cisco.
LovEjoY, T. E. 1978. Late twentieth-century con-
servation: the science of a simplifying biosphere.
First International Conference of Reserve in
Conservation Biology, Univ. California, San
Diego.
Nations, J. D. 1979. Cattle, cash, food, and forest: the
destruction of the American tropics and the La-
candon Maya alternative. Culture and Agricul-
ture, Univ. California, Davis (in press).
Parsons, J. J. 1976. Forest to pasture: development or
destruction? Revista de Biologia Tropical, Costa
Rica. 24(1): 121-138.
RiBEiRO, D. 1971. The Americas and civilization. Geo.
Allen & Unwin, Ltd., London (translated from
Portuguese).
Stevenson, W. T. 1969. History as myth/ the import for
contemporary theology. Seabury Press, new York.
Wallerstein, I. 1974. The modern world-system: capi-
talist agriculture and the origins of the European
world-economy in the sixteenth century. Aca-
demic Press, New York.
Wasserstrom, R. F. 1977. Land and labor in central
Chiapas: a regional analysis. Development and
Change. 4:441-46.3.
1978. Population growth and economic devel-
opment in Chiapas, 1524-1975. Human Ecology.
6:127-144.
Whitehead, A. N. 1933 (1955). Adventures of ideas.
Mentor Books.
PERSPECTIVE
John L. Spiiiks'
Abstract.— The fact that mankind has desecrated much of the natural world is recognized. The rate of plant and
animal extinction has increased in North America from an estimated 3 species per century 3,000 years ago to an
average of 143 per century since 1620. Endangered species protection began in the Fish and Wildlife Service in 19.38
with the purpose of the Aransas National Wildlife Refuge for the whooping crane. A committee on rare and endan-
gered species was formed in 1962 by the director of the Fish and Wildlife Service and a tentative list was published
in 1964. The Endangered Species Acts of 1966, 1969, and 1973, together with subsequent amendments, provide the
legislative authority for the present program. The intent of Congress, through this legislative authority, is to avoid
irreversible or irretrievable commitments of resources by identifying problems of environmental impact projects
early in the planning stage. Examples in the step-by-step development of the legislation and its operation were re-
viewed.
I certainly sympathize with the difficulty
that Tom Lovejoy and Roland Clement had
with their presentations prior to mine, but
with all due respect I think perspectives are
a bit difficult to address. Perspectives are
very individualistic things held certainly very
precious to those individuals who have them.
When organizations or groups have a similar
perspective on something, they're often in-
stitutionalized. I would not be so presump-
tive as to try to imply that the Fish and
Wildlife Service collectively or myself indi-
vidually has the only perspective on endan-
gered .species and endangered species pro-
grams. All we can do is hope that a general
public interest and a realistic perspective can
be gained by all of those who may affect or
be affected by our administering the Endan-
gered Species Act of 1973 as amended.
To have any perspective I think you must
have a little historical sense as to how we got
here from there. Then I want to get into the
nitty-gritty things that are not so much per-
spective as they are pragmatic problems
we're going to have in administering the
1978 amendments. We do not have all the
answers to a number of rather weighty ques-
tions presented by tho.se amendments, but I
would like you to leave here today with at
least as much knowledge as we have as to
how we're going to proceed.
The fact that we have desecrated much of
the natural world is almost given at this
point. There have been various ways to quan-
tify this. Nobody is sure what the quan-
tification means. We are not exact in saying
that it means a certain loss to us by having
made a given species extinct. At least we do
know what happened here in North America.
In the 3,000-year period prior to our arrival,
the natural extinction rate was about 3 spe-
cies per 100 years. Since the Puritans arrived
at Plymouth Rock in 1620, over 500 species
and subspecies of North American flora and
fauna have become extinct. Norman Myers
expresses the impact we have had on re-
sources, on species and subspecies in an ex-
cellent statement, condensing earth's exist-
ence down to one calendar year, as follows:
To condense the evolution of life on earth into a more
comprehensible frame of reference, suppose the whole
history of the planet is contained within a single year.
The conditions suitable for life did not develop until late
June. The oldest known fossils are living creatures about
mid-October and life is abundant for both animals and
plants, mostly in the seas, by the end of that month. In
mid-December dinosaurs and other reptiles dominate
the scene. Mammals appear in large numbers only a
little before Christmas. On New Year's Eve at about five
minutes to midnight, man emerges. Of these five min-
utes of man's existence, recorded history represents
about the time the clock takes to strike midnight.
The period since 1600 A.D., the one refer-
enced earlier, when man-induced extinction
began to increase rapidly, amounts to about
three seconds. The quarter-century just be-
gun, when the disappearance of species is put
on the scale of all the mass extinctions of the
'Chief, Office of Endangered Spec.es, U.S. Fish and WildUfe Service, Washington. D.C. 20240.
17
18
Great Basin Naturalist Memoirs
No. 3
past put together, will take one-sixth of a sec-
ond. So the process by which species have
become extinct has been incredibly accelera-
ted by the impact of man.
We have classic cases here in North Amer-
ica, such as the passenger pigeon, which once
numbered in the billions and became extinct
in 1914. It is very difficult to say what the re-
action of the people who lived in that time
might have been as these species went by the
boards. There were certainly some who were
economically sensitive of the loss. Passenger
pigeons made great feed for hogs. They could
be caught on their roost and killed by the
barrel loads with sticks, so there was some re-
action, but it was not really a societal at-
tempt, with money behind the movement, to
do something about endangered species. It
was not until the 1930s, when the 1932 Ani-
mal Damage Control Act was passed by Con-
gress (which is still in the operative legisla-
tion, incidentally, for federal activities in ani-
mal damage control), that there was a hue
and cry from the Society of Mammalogists
about consequences to vulnerable species.
Dr. E. Raymond Hall still remembers vividly
his concern as a young man for what had
happened to the gray wolf, and he did not
like the future prospects.
The Fish and Wildlife Service really began
"endangered species" protection, in terms of
major fiscal outlays, in 1938 with the pur-
chase of the Aransas National Wildlife Ref-
uge for Whooping Cranes. The cranes at that
time had reached a low of 14 birds and were
in a very critical situation. The service con-
tinued to work on whooping cranes, and in
1956 a coordinating committee was estab-
lished between the service and representa-
tives of the National Audubon Society to see
what could be done about a concentrated ef-
fort to save the whooping crane. Since then
progress has moved steadily in terms of sensi-
tivity and concern for vanishing animals, but
I would emphasize that the early concern
was more for animals and more specifically
mammals and birds than any consideration of
lesser lifeforms. If it had big brown eyes and
was cuddly or in some way looked noble,
then folks had an increased tendency to love
it and be concerned if it was disappearing.
Skuas and invertebrates really didn't turn
folks on too much then and, as a matter of
fact, they don't turn folks on very much now.
That's another story.
In 1962, a committee on rare and endan-
gered wildlife species, composed of the vari-
ous divisions of the Fish and Wildlife Service,
was formed by the director to begin wres-
tling with the problem of what should be
done with these critters for which we should
be responsible. By January 1964, a tentative
list of endangered species was put together
by the service and circulated for review, and
this resulted in 1966 as "the red book," the
good old red book you may have seen in your
libraries on native, rare, and endangered spe-
cies.
Perspectives. How did we get from the last
passenger pigeon in 1914 to a federal action
in the late 1960s? It's difficult to say. Endan-
gered species are very difficult animals to
think about and the legislation that protects
them is a very difficult type of legislation to
understand. I think one perspective that folks
have on the Endangered Species Act reminds
me of Mark Twain's comment on the Bible.
He said that he didn't understand very much
of it, but what he did understand scared the
hell out of him. In many respects this is
where we have been with endangered species
legislation. The first Endangered Species Act
of 1966 was a rather innocuous piece of legis-
lation, in all honesty, and particularly so
when compared to the 1973 act. It allowed
us to list native, endangered species and to
acquire land with Land and Water Con-
servation Fund monies. There was no pro-
cedural requirement as to how things were
put on the list, however, and it didn't do a
critter a lot of good because being listed af-
forded no protection from taking. It was,
however, a first fledgling step to a mean-
ingful national law protecting endangered
species. Also, there was only one category, an
endangered species, and an endangered spe-
cies was basically a basket case, something
that was in dire straits. Reference was made
to rare species in the red books published in
1966 and 1968 but rare species were not in-
cluded in legislation. In 1969, a second en-
dangered species act was passed. The Endan-
gered Species Conservation Act, and this act
went a bit hirther than the 1966 act. It did
broaden the definition of fish and wildlife to
include moUusks and crustaceans, a rather
1979
The Endangered Species: A Symposium
19
large step forward because, heretofore, pre-
dominant concern had been with vertebrates,
mostly those that were lovable. The Lacey
Act was amended to allow a broader degree
of enforcement by including reptiles, am-
phibians, moUusks, and crustaceans. Foreign
species could be listed for the first time under
the 1969 act. A very important international
step was taken by the 1969 act when the sec-
retary of the interior was directed to seek the
convening of an international ministerial
meeting before 30 June 1971, at which would
be concluded a binding international conven-
tion on the conservation of endangered spe-
cies. That convention took place and is now
the Convention on International Trade in
Endangered Species or Wild Fauna and
Flora, a very important international agree-
ment to which some 46 countries are now
parties. Then came the big one, the Endan-
gered Species Act of 1973, which President
Nixon signed into law on 28 December. The
Endangered Species Act of 1973 could accu-
rately be described as a "sleeper." I am sure
Congress was unaware of the fviU implica-
tions of its provisions.
Tellico Dam is a good case in point. Tell-
ico had been imder litigation from local citi-
zens who were opposed to it for a number of
years before the snail darter swam into the
picture. Perhaps Tellico and the snail darter
could be likened to the whale who swallowed
Jonah under inverse circumstances. Jonah
swallowed the whale and the snail darter
seems to have engulfed Tellico Dam. After
the snail darter was scientifically described,
an emergency rule making listed the species
and determined its critical habitat. We were
petitioned to do so.
The federal district court in which the case
was first tried did not find for the plaintiffs.
In addressing the issue of saving either Tell-
ico Dam or the snail darter, they found for
the Tennessee Valley Authority. The district
court's decision was appealed. It was re-
versed in the federal appellant court and ulti-
mately came before the U.S. Supreme Court.
The Supreme Court also ruled for the snail
darter but not in the true context of that
statement, in that the Supreme Court said,
"Yes, this is really what the Endangered Spe-
cies Act says. This is what Section 7 of the
Endangered Species Act says." It says that all
federal agencies shall insure that actions au-
thorized, funded, and carried out by them do
not jeopardize the continued existence of a
threatened or endangered species or adverse-
ly modify or destroy its critical habitat, and
that's what the TVA's actions were clearly
going to do. There was no question of that
being the ultimate outcome should the dam
be completed and the gates closed.
The ripple that reached tidal wave propor-
tions following the decision could perhaps be
characterized as the "Chicken Little Syn-
drome." Do you remember Chicken Little?
Chicken Little was out in the barnyard one
day when an acorn dropped on his head and
he assumed the sky was falling. Other parties
with similar federal works projects saw the
acorn fall on Tennessee Valley Authority's
head and assumed the sky was going to fall.
It was Chicken Little all over again. There
was a deep concern that economic progress,
if you will, inckiding many important public
works projects, would be halted because of
endangered or threatened species being pres-
ent.
We felt in the Fish and Wildlife Service at
that time, and we still do to this day, that the
concern was an overconcern, that we could
find no justification for it. The service had
completed some 5,000 formal and informal
consultations with other federal agencies.
Three of those at that time had been liti-
gated. In one instance involving the Indiana
Bat and Merramac Park Lake, the court
found in favor of the U.S. Army Corps of En-
gineers both at the district court level and
the appellant court level. In the second case
involving the Mississippi Sandhill Crane and
Interstate 10, the court did find for the
plaintiffs, but that highway has since been
completed. The questionable interchange is
going in. The conflict was resolved ulti-
mately by the Fish and Wildlife Service and
the Federal Highway Administration work-
ing cooperatively, so we could have our cake
and eat it, too— or have our cranes and their
interchange, too, if you want to put it like
that. We felt there was a degree of over-
reaction to the problems that were going to
be caused by Tellico. We thought it was an
anomaly. It was not typical of what the En-
dangered Species Act was going to do in the
future. Nevertheless, a number of individuals
20
Great Basin Naturalist Memoirs
No. 3
were concerned about this, and a number of
bills were put before Congress to address the
problem. There were any number of varia-
tions on these bills, including specific exemp-
tions for the Tellico Dam and for another
TVA project on the Duck River. Some 14 or
15 bills were being considered by Congress,
some introduced in the House of Representa-
tives, some in the Senate.
The first thing that happened, in terms of
action, was a Senate bill, cosponsored by Sen-
ator Culver of Iowa and Senator Baker from
Tennessee. This bill presented a mechanism
by which an appeal could be made and a
project exempted from the Endangered Spe-
cies Act. A focus was finally made in the
House of Representatives on a bill reported
out of Mr. Legget's subcommittee which had
that provision as well as a preliminary review
step by a review board. The outcome of all
this was an amendment to the Endangered
Species Act which passed Congress in the
eleventh hour on 14 October, just before
Congress was going to adjourn. Unfortu-
nately, our appropriation authority to admin-
ister the Endangered Species Act had expired
at midnight on 30 September. We were out
of business for two weeks because we had no
money to operate the program. The act itself
remained in effect, the prohibitions of the act
remained in effect, and our obligations to
consult remained in effect. However, we had
no money to do any of these things. President
Carter signed that bill on 10 November at
10:00 p.m. That was the last day the presi-
dent had to sign the bill before it was pocket
vetoed. That made a total of 41 days that the
Office of Endangered Species, indeed the en-
dangered species program, was out of busi-
ness.
We are now back in business. We're dig-
ging out and we're trying to understand the
1978 amendments to the Endangered Species
Act. I want to go over these with you briefly.
They are too complex to focus on in great de-
tail. Again I would qualify an ultimate con-
sideration of what these amendments say to
the extent that, until we have a firm reading
from our solicitor's office on some finer inter-
pretations of the intent of Congress, we are
going to be walking a tightrope blindfolded
at times to try to administer these and keep
the intent of Congress uppermost in our
minds.
One of the more interesting happenings
was a redefinition of critical habitat. There
had been no definition of critical habitat in
the original 1973 act. It was mentioned in
Section 7 of that act and it was defined by
regulation by the Fish and Wildlife Service
in the Section 7 regulations. The new defini-
tion basically confines critical habitat to the
geographical area in which a species present-
ly occurs. It does make allowance for consid-
eration of specific areas outside the geo-
graphical area where the species is found, but
only if these areas are determined to be es-
sential for conservation. What does essential
for conservation mean? Conservation is de-
fined in one place in the act, but we are not
sure what the degree of essential is.
Another important happening was the def-
inition of species. Tom Lovejoy alluded to
the lessening of protection for invertebrates
and, at one point, in one earlier bill which
was not enacted, there was a rather glaring
distinction made against invertebrates— as I
recall, something to the effect that they
could not have critical habitat determined
for them. That was changed in the final act.
The major difference made between in-
vertebrates and vertebrates is that we cannot
list invertebrates at the population level.
They may only be listed at a subspecific lev-
el.
Now for Section 7 itself. The key elements
for requiring an agency to consult with the
Fish and Wildlife Service, if their activities
may affect a listed species, are still in place.
This has not changed at all. As a matter of
fact, the necessity for consultation has been
stengthened by these amendments because,
without a good-faith consultation, an agency
will not qualify for an exemption under other
provisions of the act. There is more definition
given to the opinion to be rendered by the
secretary of the interior, i.e., the director of
the Fish and Wildlife service to whom the
authority to administer the act has been dele-
gated. It now specifies what must be con-
tained in the biological opinion.
An entirely new element called a biologi-
cal assessment has been introduced which
only applies to agency action for which no
contract for constniction has been entered
1979
The Endangered Species: A Symposium
21
into and for which no construction has begun
on the date of the amendments. A biological
assessment must be done on projects that fall
in this category. The agency that has the ac-
tion must request from the secretary of the
interior a list of proposed or listed species
which may be found in the project area. The
agency has 180 days in which to conclude a
biological assessment to see what indeed is
there. The intent of Congress is that you find
out the problem in the early planning stage
before you get in the middle of a dam and
then end up with another confrontation on
your hands. During this process and during
the consultation process, the action agency
cannot make an irreversible commitment of
resources.
A federal agency, the governor of a state in
which a project is located, or a license or
permit applicant whose permit or license is
being denied because of endangered or
threatened species can appeal for exemption
to an endangered species committee. The ap-
pellant has 90 days after a biological opinion
has been rendered in which to submit this ap-
peal. The endangered species committee is
composed of seven members, the chairman of
which is the secretary of the interior and the
other members being secretaries of agricul-
ture and the army, the chairman of tlie Coun-
cil of Economic Advisors, the EPA, the ad-
ministrator of NOAH, and one person or
persons appointed by the president from the
state(s) affected by the project action.
Before the committee gets to look at the
exemption or the request for one, however,
it is first referred to a review board, a sec-
ond-tier process which was not included in
the Baker-Culver amendment from the Sen-
ate. This review board has three persons on
it, one appointed by the secretary of the inte-
rior not later than 15 days after the appli-
cation, one appointed by the president, and
an administrative law judge. It is the job of
this review board to examine the application
for exemption, and they look at four basic fac-
tors: (1) Does an unresolvable conflict exist?
(2) Has the agency carried out the con-
sultation in good faith? (3) Did it conduct the
biological assessment required of it? (4) Did it
refrain from making an irreversible com-
mitment of resources?
Within 60 days after receiving the appli-
cation for exemption, the review board must
have been appointed and have positively de-
termined that these criteria have been met.
The board reports to the committee, and
within 180 days after they make a determina-
tion they must recommend to the committee
reasonable and prudent alternatives to the
action, summarize the evidence as to whether
or not the agency action is within the public
interest and of national or local significance,
and decide if mitigation and enhancement
measures should be considered by the com-
mittee. Once the committee gets all this in
hand it has 90 days to decide whether or not
it will exempt a project from the require-
ments of Section 7. In the process of doing
this, the committee must make four findings:
that there are no reasonable or prudent alter-
natives to the agency action, that the benefits
of the action clearly outweigh alternative
courses consistent with preserving the species
or its critical habitat, that such action is in
the public interest, and that the action is of
regional or national significance. However,
after proceeding this far in the exemption
process, if the secretary of the interior deter-
mines the exemption would cause the extinc-
tion of a species, he so advises the committee
and the committee has 30 more days in
which to decide whether or not the project
will cause the extinction of a species by vir-
tue of granting an exemption to the agency
action. There is also a review provision by
the secretary of state that if the exemption
would violate any international treaty or ob-
ligation then the exemption cannot be al-
lowed. This will be addressed in the regu-
lations promulgated by the committee itself.
They have 90 days after enactment of the
1978 amendments to propose these regu-
lations.
This is the core of how the exemption pro-
cess works, only the core. The complete, re-
vised version of the act, with the 1978
amendments incorporated, will be available
from the Fish and Wildlife Service sometime
around 1 January. At the present time, we
only have a copy of the signed bill itself and
this can be rather confusing unless you are fa-
miliar with the 1973 act and can see where
all the "wherefore's" and "thou art's" go.
One other thing that the amendments did
was to provide for immediate consideration
22
Great Basin Naturalist Memoirs
No. 3
of exemption for both Tellico Dam and Gray
Rocks Dam. The committee has 30 days to
begin consideration of both projects and 90
days to decide whether it will exempt. If it
fails to act within 90 days, both projects are
exempted by virtue of this statute.
There was an amendment to Section 6 of
the act which for the first time brings plants
under the purview of the grant-in-aid pro-
gram. Heretofore Section 6 cooperative funds
were only available for animals, not plants.
Also, the bill authorized our expenditures un-
der the act. I indicated earlier that we went
out of business when our appropriation au-
thority expired. We only received 18 months
of reauthorization, which means we will go
through the same process of having the act
reauthorized in 18 months. We anticipate
oversight hearings on the Endangered Spe-
cies Act this spring, probably in both houses
of Congress.
What we are going to do about getting on
with listing of endangered and threatened
species and determining critical habitat for
these species is something else again. We had
originally planned on some 200 rulemakings
in fiscal year 1979. Our present estimate is
that maybe 20 to 30 rulemakings will be pos-
sible. The reason for this is the greatly in-
creased workload to list a species. It will be a
more expensive process; it will be a more
time-consuming process. Some of the ele-
ments involved in the new listing process are
good: holding public hearings, notifying local
people that an action is contemplated, pub-
lishing in a local newspaper. We think that
the increased public involvement in the deci-
sion-making process will be beneficial in the
long run.
We hope we can resolve some of the con-
cerns that have been expressed over many
proposals. It appears, however, that there are
a couple of "Catch 22's" in terms of present
proposals. There is a two-year expiration pro-
vision in the 1978 amendments. It says, in ef-
fect, that, if a species or critical habitat has
been proposed for two years and it hasn't
been finalized within that two-year period, it
expires and must be withdrawn by the secre-
tary of the interior. There is a one-year grace
period, however, for existing proposals. That
one year will be up on 10 November 1979.
Over 1,700 plants are proposed. We realize
we will be able to list perhaps a fraction of
those. All of the existing critical habitat pro-
posals will more than likely be withdrawn
because of the new requirements involved in
determining critical habitat. Those require-
ments include doing an economic analysis
and an analysis of other relevant impacts and
we're not sure what other relevant impacts
really means. Here again the lawyer will
come to our rescue.
We are going to place in priority form the
existing proposals based on degree of threat
before the on-year expiration period comes
up. We do not have a large staff in the pro-
gram. Basically the law charges us with the
responsibility for the animal and plant king-
doms of the world. We have something less
than 200 permanent full-time positions with-
in the endangered species program split be-
tween the Office of Endangered Species,
Federal Wildlife Permit Office, the Division
of Law Enforcement, and the National Wild-
life Refuge System. So the dilution of per-
sonnel across the program scope is tre-
mendous. It is a challenge, a challenge which
we welcome, and the espirit de corps within
the program has never been higher.
Back to perspectives again. Perspectives
are very difficult. At times it is difficult to
justify, depending on the individual's per-
spective, listing a species and perhaps imped-
ing a given project. The question keeps com-
ing back. What good are endangered species
or threatened species? Tell us in a very tan-
gible fashion what good a snail darter is. We
cannot answer that. We cannot give you a
dollar and cent answer to that kind of ques-
tion. The most lucid comment which address-
es this concept, however, is one which was
made by Aldo Leopold, who said that the
first sign of intelligent tinkering is that you
don't throw away any of the parts. With all
of our sophistication, I think we are tinkering
with phenomena that are much more sophis-
ticated than we. Ovir concern is certainly for
the survival of the species. It is also for the
survival and well-being of mankind. It is our
posture that, until our knowledge, as a race,
as a society, evolves to the point that we can
clearly know the consequences of our action
by making a species extinct, it is very, very
foolish to do so. It may be the part that we
needed to make the clock run for another
centurv or so.
1979
The Endangered Species: A Symposium
23
Qu
ESTIONS TO
Dr. Si
Q. Have I been given an impossible task then to provide
for the Fish and Wildhfe Service in Utah the data on
200 endangered species of Utah plants?
A. If you think we're going to do it next week, you'll be
disappointed. If you think we're not going to do it at
all, you're wrong. It's going to be a lot of jumping
through hoops. We've had some other difficult
hoops to jump through and our intent is to get this
program unwound as rapidly as we can. We've been
digging out from 41 days of inactivity, but I feel
rather confident in telling you that your data is not
going to be gathering dust for an indefinite period of
time. If those species for which you are providing
the information fall out as priority species having
the most danger, the greatest degree of threat,
they'll be among the first we get to.
Q. Is there any aspect of litigation involved in this new
amendment? In other words, how do we give people
the chance to question something we say is becom-
ing extinct, like the snail darter? Does the applying
agency have to provide research fimds or try to relo-
cate the snail darter even though they might not be
successful in that aspect of threatened or endan-
gered species?
A. Yes. The committee will actually direct the appeal-
ing agency as to what must be done on behalf of that
species, and the agency taking the action is respon-
sible for bearing the cost of that. Now in terms of
construction projects, this cost is not considered in
evaluating the cost-benefit ratio of the project. It
will be an additional cost; but it would not, for in-
stance, bring a project below parity and thereby
make it unfeasible or illegal to build.
Q. In all the time limits that have been set, the 30 days,
the 90 days, the 180 days, what happens if an agency
or committee fails to meet these deadlines?
A. There is no slap on the wrist if anyone fails to meet
the actual time frame. Some of those time frames,
incidentally, are negotiable in that the 180-day bio-
logical assessment could be lengthened if the agency
requested it with agreement between the agency
and secretary. If you add up all the maximum time
frames, however, including the 180 days, the total is
something like 750 days that the entire process could
take.
Q. But there is no traditional mechanism?
A. No, but the citizen suit provision of the act still ap-
plies, and anyone could litigate against any party
that failed to meet its deadlines.
Q. It has been the thmst of the whole program all along
that the brunt of the responsibility has fallen on oth-
er federal agencies, besides the Fish and Wildlife
Service, and private organizations, too. But isn't it
true in the West, where field work for proposed spe-
cies is just starting? Now suddenly I'm being pushed.
I know I'm speaking to you in a sense, but I'm also
speaking to me. I'm one who elected the people who
are passing these things, but 20 or 30 are not going
to be enough. We need more people. There are a lot
of areas where work needs to be done.
A. There is a "Catch 22" in everything, I guess. There's
also a hiring freeze in the federal government at the
moment which affects permanent, full-time posi-
tions. .\s a matter of fact, there is nothing we can do
about that. 1 hope you can also appreciate the diffi-
culty of bringing in a permanent part-time or some
other less than permanent position and expecting
that person to walk in and start doing something
productive the next day. It takes a lot of expertise
and training to write a decent rulemaking, for in-
stance, one that will get by the scrutiny of the solic-
itors and be legally justifiable and adequate.
Q. .\11 I'm asking is to just make an effort.
.\. We are.
Q. Pertaining to the exemption process, other than liti-
gation, where is the avenue for public involvement?
A. There is a provision which provides the meetings of
both the review board and the committee to be
open. It will depend on whether the committee de-
cides to take testimony from the public. That point,
I'm sure, will be addressed in regulation pro-
mulgated by the committee and by the review
board. The final decision of the committee is subject
to judicial review. It can be appealed to the courts,
and there is specific provision in the legislation for
that.
Q. When would you determine the rulemaking for criti-
cal habitat for the grizzly bear?
A. As I indicated earlier, it is very likely that all exist-
ing proposed mlemakings for critical habitat will be
withdrawn. In effect, that proposed rulemaking
would be invalidated and a reproposal would come
forth. The reproposal would have to meet the new
criteria of the 1978 amendments, including an eco-
nomic impact analysis and identifying actions or ac-
tivities within the area, which might be affected by
having the area designated as critical habitat— both
federal actions as well as private actions. We do not
have an economist on our staff and, quite frankly, it
gives us .some heartburn to consider a meaningful
economic analysis. I am not being facetious when I
say meaningfid, because we're not going to try to
short-cut the intent of Congress in this thing. They
want an economic analysis, one that is meaningful,
and that is what they are going to get from us. We
don't know where the help is going to come from,
perhaps from within the department and other
agencies which do have economic expertise.
A. You recently listed some species in California with-
out listing critical habitat. Are these being consid-
ered for withdrawal under new amendments?
.\. No, anvthing that is already listed that did not have
critical habitat determined at the time it was listed
will remain a listed species. The amendments say
that we may determine critical habitat for these spe-
cies at some point in time. We can do this; we don't
have to do it yesterday. What we do have to do in
the future, however, unless it is pnident not to do so,
is to propose critical habitat at the same time we
propose listing of species, so these two things go
along simultaneously. There was no provision for
critical habitat in either the 1966 or 1969 acts. That
is why we have a huge backlog of listed species that
have no critical habitat.
Q. Isn't it true that any agency must consult the Fish
and Wildlife Service before beginning any project?
24
Great Basin Naturalist Memoirs No. 3
Do we mean any project or are we defining proj-
ects?
No, when an agency identifies that its actions may
affect listed species, that is when they must initiate
consultation. It is the may affect. Now the confusing
element here may have been my comments con-
cerning constmction contracts, projects for which no
contracts have been let and for which no constmc-
tion has begim. These are the ones that would have
to do a biological assessment before things could
proceed if there are listed or proposed species in the
area, but that is different than consultation per se.
THE LAW AND ITS ECONOMIC IMPACT
Donald A. Spencer'
Abstract.— There is no adecjiiate inventory of population size and distribution of most of the world's animal and
plant species and lower taxa. Furthermore, populations are rarely static and continue to change in response to both
natural and man-made factors. Thus clearance today for public works or industrial projects can be reversed tomor-
row as new information becomes available. Lacking assurance that a project can be completed without new endan-
gered species surfacing places an untenable constraint on the commitment of dollars for new long-term programs.
As a consequence of the absence of data, studies to determine occupied range, population levels, and habitat re-
quirements of specific endangered species must be conducted on each project area. The direct costs of these studies
are the responsibility of the project applicant. The time consumed results in project delays which can become a
major expense item. Additional economic impacts are inherent in construction modifications and subsequent project
operations intended to accommodate an endangered species.
Finally, the withdrawal of natural resouces to support endangered species can conceivably reach a point where
the squeeze on other societal programs becomes unacceptable.
Thank you for making this time available
to me. It's always a privilege to get together
with a group that is intentionally interested
in a good program and talk problems out.
I brought along this book. I thought some
of you might want to obtain a copy of it. It is
a proposed environmental impact statement
on the effect of grazing on some of our west-
ern lands. This little book cost $250,000 to
prepare. It is, actually, an excellent study;
you'll be much impressed by what the au-
thors and the various research teams have put
together in it. But as I read through it and
came to the areas of my own expertise, I
found that, if we are going to consider endan-
gered species on this 800-square-mile area
this book is about, we're going to have to do
the job. The information on endangered spe-
cies, on wildlife and nongame species in gen-
eral, is treated once over lightly.
In the back of the publication, I began to
read the letters received about this program
from people who actually lived on the area;
and who were going to be affected by it, not
an outsider like me who was reading what I
considered to be a very excellent program.
Then it occurred to me that this was just like
what had been happening to me. You know
most of my professional career has been
spent in research in the field of animal biolo-
gy. When you come up with a new tool or a
new program or a new project that is the re-
sult of research and you're very proud of ac-
complishing something new for wildlife man-
agement, you send the report all over the
coimtry for trial. When it comes back to you
from first one point then another, reviewers
state that it won't work here or that it pro-
duces an adverse affect there. You're very
bitter about it— you even tend to react vio-
lently. Then you begin to realize that of all
tlie things research values most highly it is
knowing the limitation of the new tool.
Where are the boundary lines where it works
most effectively? If you do not recognize
those boundaries early, you are liable to lose
the use of the tool in the areas where it
would be valuable. There are literally hun-
dieds of examples in the last four or five
years to bear this out and I won't have to
elaborate.
One thing we have to realize is that in ask-
ing for habitat for endangered species we are
in competition, and I'm using that term ad-
visedly, with a lot of other conservation ob-
jectives. For example, over 100 years ago we
began a national park system that has grown
to some 300 units encompassing in excess of
31 million acres. The Brown Pelican, which
has been so much in the news of recent years,
initiated the first unit of the National Wild-
life Refuge System that has now grown to 34
million acres. We have now reserved be-
tween 100 and 150 million acres where wild-
'13508 Sherwood Forest Terrace, Silver Spring, Maryland 20904.
25
26
Great Basin Naturalist Memoirs
No. 3
life receives top billing.
Building on these established programs,
and adding such new ones as the Wild and
Scenic Rivers (1968), National Sea and Lake
Shores, Wilderness (1964), National Trails
System, Marine Protection Research and Es-
tuaries (1972), Research Natural Areas,
Coastal Zone Management Act, Agriculture's
Water Bank Program, Wetlands, etc., has be-
come a veritible national obsession. Cur-
rently President Carter, under the authority
of the Antiquities Act of 1906, has pro-
claimed 56 million acres in Alaska as national
monuments, and the secretary of the interior
has temporarily withdrawn an additional 54
million acres from any commercial devel-
opment. The U.S. Forest Service has under
review some 62 million acres of "roadless
areas" for possible inclusion in the Wilder-
ness System. The Bureau of Land Manage-
ment, overseer of 470 million acres of public
lands, is engaged in a similar "roadless area"
review to determine what lands of theirs
would qualify for wilderness designation.
In a very limited way, habitats purchased
specifically to protect an endangered species
add to the set-aside totals. But the greatest
impact will result from "critical habitat" des-
ignations that, while not infringing on any
use that does not adversely modify the habi-
tat for a given endangered species, still im-
pose costly constraints on change, to the
point of completely preventing some projects
useful to man. Alternate uses that are affect-
ed by critical habitat constraints can be quite
varied, as the following from the U.S. Forest
Service's Wildlife Management Manual illus-
trates:
Maiiv projects and practices authorized or carried out
hv the Forest Service are of such a nature that modifica-
tion of the vegetation or land is often a direct or indirect
result. These include activities such as recreation site
development, land exchanges, timber sales, revegetation
and reforestation, type conversions, water impound-
ments, road and trail construction, grazing by herbi-
vores, and development that results in significant in-
creases in the level of human activity in an area.
The patterns for establishing critical habitats
give little as.surance that very many areas
will be free of constraints to protect one or
more endangered plants or animals. A num-
ber of critical habitats, both established and
proposed, are disturbingly large. The critical
habitat for the Manatee includes every major
estuary in peninsular Florida, sits astride the
busy intercoastal waterway, and is one of the
most intensively used recreational boat areas
in the United States. The proposed critical
habitat for the grizzly bear suggests 13 mil-
lion acres in Montana, Idaho, and Wyoming
which encompasses two national parks, Yel-
lowstone and Glacier. Throughout the visitor
season, a succession of trails, campgrounds,
and back-country are closed to people be-
cause of bears. As for the Whooping Crane, a
proposal will about triple its present 90,000-
acre wintering ground and provide seventeen
migratory stopovers in six states and a new
nonhistorical experimental breeding range in
three other states. Some of these brief stop-
over points are not necessarily small areas.
Along the Platte River in Nebraska the linear
54 miles of bottom land totals about 103,000
acres; the proposed migratory stopover on
the Niobrara River is 115,200 acres; the large
proposed area along the Canadian border in
northwest North Dakota probably exceeds 2
million acres. Then, surprisingly, it is pro-
posed to include the dams and lake margins
between maximum and minimum pool of the
two largest flood control impoundments on
the Missouri River (Lake Oahe and Lake
Sakakawea).
As tne endangered list grows by the addi-
tion of relatively little-known species and
subspecies from the enormously large pool of
living plants and animals these resource set-
asides can provoke a reaction that will dam-
age even the best features of the program.
The Regulatory Thicket
The independent, consumer-owned power
companies serving some 200 municipalities in
eight Missouri Basin States found their pres-
ent capacity for electric power desperately
below what would be needed in the years just
ahead. So they formed the Missouri Basin
Power Project, which began constructing its
first generating plant, the Leland Olds Sta-
tion, in 1962. The construction required only
one government permit and was completed
in four years. Unit No. 2 at this site required
five permits and went on line in 1975. Their
next cooperative project, the Laramie River
Station and Grayrocks Reservoir, got under
construction in 1976 and has thus far re-
1979
The Endangered Species: A Symposium
27
cjuiied 43 pennits and approvals from feder-
al, state, and local authorities. Their third
project, the Antelope Valley Station, begun
in June of this year (1978), has at this early
stage required 69 federal, state, and local per-
mits—a\\ of this within the experience of one
vital cooperative project serving an eight-
state area.
Like illustrations of the maze of govern-
ment regulations are at every hand. For ex-
ample, the atomic-powered electric plant un-
der construction at Midland, Michigan, has
acquired some 93 pennits to date.
The redundant nature of some of these
regulations is a sad commentary on the effi-
ciency that has marked so much of America's
progress. Take, for example, the construction
of a transmission line that links North Dakota
and Minnesota. Not only did the federal Ru-
ral Electrification Agency require an Envi-
ronmental Impact Statement (Study), but a
Certification of Need had to be obtained
from the Minnesota Energy Agency, a Cer-
tificate of Corridor Compatability from the
Minnesota Environmental Quality Board, and
two separate permits for Site Compatability
and for Route Designation from the North
Dakota Public Service Commission. The
transmission corridor crossed four navigable
rivers requiring four separate construction
permits from the U.S. Corps of Engineers. It
made eleven other water crossings, each re-
quiring separate permits from the Minnesota
Department of Natural Resources; crossed
three wetland areas requiring as many ease-
ments from the U.S. Fish and Wildlife Ser-
vice; made eleven highway crossings, each
requiring a permit from the respective coun-
ty highway department— which was in addi-
tion to two separate permits from the North
Dakota Land Department, one labeled State
Land Crossing, the other River Crossing; and
crossed five different railroads requiring per-
mits for each location. It can be agreed that
such a corridor has to obtain easements
across every privately owned parcel of land,
so why should public land be different? The
point to be made here is the large number of
governmental agencies have replicated input
into a single project.
A review of Sections 7 Consultation Logs
of the U.S. Fish and Wildlife Service's six re-
gional offices for the period of October 1977
through May 1978 reveals that between 30
and 35 different federal and state agencies
contacted the Office of Endangered Species
for advice on their responsibilities under the
act.
With such a galaxy of regulatory agencies
afield, there are few, if any, projects that do
not require a permit or license of some kind.
A western cattleman will need a grazing per-
mit to use public land, a farmer will need a
point source discharge permit for return irri-
gation flows, to build a dock or bulkhead on
your waterfront property will require a per-
mit, and, even if you wish to participate in
the recovery of a Peregrine Falcon, thus en-
hancing this endangered species, you must
have a permit. Regulation reaches into the
most remote corner of our society.
The regulatory morass motivated President
Carter in March 1978 to issue Executive Or-
der 12044 "as a first step toward ensuring
that regulations achieve their statutory goals
in the most effective and balanced way."
This now has been followed up by the Presi-
dential appointment of a Regulatory Council
to inform me, the public, and the Congress about the
cinmilative impact of regulation on the economy. The
Council will help ensure that regulations are well
coordinated, do not conflict, and do not impose excess
burdens on particular sectors of the economy. The first
report of this Council is to be made public no later
than February 1, 1979.
At reoccurring intervals officials of the Office
of Endangered Species have sought to clarify
what is meant by critical habitat by saying
that the act charges federal agencies— and
only federal agencies— with carrying out pro-
visions of Section 7. State and private actions
not involving federal approval do not come
imder the act. This is meaningless comfort
when it would be almost impossible to identi-
fy a private project that does not need some
federal (or state) approval. Even if this were
not so. Section 9 provides severe penalties for
any person who "harasses, harms, pursues,
hunts, shoots, wounds, kills, captures, col-
lects, or attempts to engage in any such con-
duct" an endangered or threatened animal no
matter where found. Thus the act, for all in-
tents and purposes, affects public works and
private projects alike.
The Permit: Often Elusive
The permit or license is elusive because it
28
Great Basin Naturalist Memoirs
No. 3
can be withdrawn by the issuing agency on
the basis of new information not previously
considered, and because a court of law can
order the permit suspended or withdrawn
pending the outcome of a public interest suit
which may rest only on some omission in the
Environmental Impact Study. It can be dis-
ruptive because the regulation, or the factor
requiring it, did not come into being until the
project was partly completed.
Advancing technology has made most proj-
ects more complex to construct. Added safety
and environment features, and the time-con-
suming efforts to comply with over-numerous
regulations, all combine to lengthen con-
stRiction time. Thus, to bring a major coal
mine to full capacity, or a new power plant
on line, can take ten years. Each year that
passes sees a 10 percent increase in construc-
tion costs. With no assurance that a project
can successfully negotiate the ever-changing
maze of regulations, financing of projects
that cannot be completed in a reasonably
short time becomes quite a gamble.
The "biological opinion" resulting from a
Sections 7 Consultation with the Office of
Endangered Species is an agency approval (if
granted)— in practice it has the force of a per-
mit. A few examples will show that these bio-
logical opinions can exhibit every one of the
above three deficiencies with respect to
clearing the project for completion. Follow-
ing completion of a broad Environmental Im-
pact Study, the Bureau of Land Management
received the following Section 7 biological
opinion on a proposed phosphate mine on
the Osceola National Forest in Northern
Florida:
It is my biological opinion, subject to the conditions
identified herein, that the proposed project is not likely
to jeopardize the continued existance of the endangered
or threatened species listed above or result in the de-
struction or adverse modification of their critical habi-
tats.
The conditions imposed are as follows:
The Bureau of Land Management must reinitiate Sec-
tion 7 Consultation should (1) new information reveal
impacts of the above-listed species or their habitats
which was not considered in this consultation, (2) the
proposed leasing [be] subsecjuently modified, or (3) a
new species [be] listed that may be affected by the pro-
posed action.
The above clearance named only species
on the established list. It did not mention that
standing in the wings waiting to go on stage
was a proposed list of plants and animals al-
most 10 times as long. Nor did it include
Florida's official state list which names still
other species. Lastly, on a project of this size,
52,000 acres, it should not be too difficult a
task to come up with an undescribed species
or one with very local distribution that has
not yet been proposed for listing.
A well-known example of the late surf-
acing of an endangered species is provided
by the Tellico Dam in Tennessee, where the
small fish had not even been described as a
distinct species of Darter until well after con-
struction had begun.
More recently the Office of Endangered
Species established a 54-mile stretch of the
Platte River bottom in Nebraska as critical
habitat for whooping cranes during migra-
tory stopovers. Upstream some 275 miles was
a power plant already under construction by
the Missouri Basin Power Project. Extraor-
dinary steps had been taken at the planning
stage of this facility to have a conservation-
acceptable project. Now, despite their hold-
ing all the required federal and state permits,
their use of water from the Laramie River
has been challenged in court because it might
reduce by 4 percent the flow of water
through the whooping crane's fall and spring
stopover. This action threatens the
$444,000,000 already invested in the project.
Each lost day will cost $140,000 in interest
on the money alone (or $50,000,000 a year).
Here, as in several other notable cases, the
Endangered Species Act is being used to ac-
complish an entirely different objective. It is
actually a matter of the continuing wrangle
over water rights between the State of Ne-
braska, who initiated the court suit, and the ■
State of Wyoming. Previously, the U.S. Su-
preme Court had awarded the water in the
Laramie River to Wyoming, so there ap-
peared to be no problem about the Gray-
rocks impoundment. What is in-
comprehensible to me is that midway
between Grayrocks and the whooping crane's
critical habitat is the Kingsley Dam in Ne-
braska that backs up Lake McConaughy,
which is 20 times the size of the incompleted
Grayrocks. The withdrawal of irrigation wa-
ter at the King.sley Dam must have marked
1979
The Endangered Species: A Symposium
29
influence on the regulated flow in the Platte
River.
The Informal Consultation
The Endangered Species Act of 1978, Sec-
tion 7(c)(a) directs—
each Federal Agency shall . . . request of the Secretary
information whether any species which is listed, or pro-
posed to be listed, may be present in the area of such
proposed action. If the Secretary advises . . . that such
species may be present, such agency shall conduct a bio-
logical assessment for the purpose of identifying any en-
dangered species which is likely to be affected by (the
project).
It is analogous to being directed to a well
for a drink, a well that hasn't been dug deep
enough to strike more than a suggestion of
water, then being drafted into the work force
to dig the well deeper. Although the act sug-
gests that this responsibility might be dis-
charged in 180 days, a review of past direc-
tives from the Office of Endangered Species
shows that such studies can take from a few
months to several years, depending on the
complexity of the biological assessment.
A number of federal agencies have elected
to prepare in advance of requests for specific
resource use, an Environmental Impact Study
covering districts or broad subdivisions. For
example, the BLM has programmed studies
on 173,919,000 acres of public lands subject
to grazing. The estimated budget to provide
the 142 separate studies at current prices is
$35,500,000, an average of $250,000 each. To
date, 16 EIS have been completed, but the
entire area will not have been covered until
1988. The proportion of this program that
can be assigned to providing data on endan-
gered species is usually quite minor. For ex-
ample, a breakdown of an Environmental
Impact Study for a proposed surface mining
project in Wyoming, which cost a private
company $500,000, shows the wild-
life/vegetational fraction costing $190,000,
with only $18,000 (or 3.6 percent) related to
endangered species information.
Another example: The Departments of
Transportation in the six lake states routinely
asked for advice if they proposed repairing or
replacing a bridge. In fact, in the period of
October 1977, through May 1978, 20.9 per-
cent of the entries on the consultation logs in
U.S. Fish and Wildlife Service, Region No. 3,
concerned bridges, and another 20.5 percent
highway actions. Just as routinely they re-
ceived the following informal "opinion":
Survey for endangered species or their habitat in the
project area, (or) If through your investigations you find
an endangered species or their habitat in the project
area you should initiate a formal con.sultation.
In most instances involving bridges the in-
terest was in one or more species of endan-
gered freshwater mussels, which required the
services of a qualified malacologist for under-
water surveys and species identification.
Aside from waiting for the proper season to
do the work, these local studies could be
completed rather quickly, as in the case of a
contract to search 49,500 square feet of the
Wabash River near Hutsonville, Illinois, at a
cost of $2,500.
The Corps of Engineers, responsible for
maintaining channel navigation in the upper
Mississippi River, finds these same endan-
gered freshwater mussel surveys far more
costly and time consuming. They have
awarded five research contracts totaling
$263,977, four of which are now completed.
A final report. Freshwater Mussels of the Up-
per Mississippi River, prepared for the corps
by the Academy of Natural Sciences of Phila-
delphia, is a 400-page document. Please un-
derstand that this required study involved
only freshwater mussels. Who is to say what
other species and subspecies of freshwater in-
vertebrates, fish, amphibians, or aquatic
plants will require similar treatment in the
future?
The magnitude of some of these studies to
determine the impact of development proj-
ects on the environment is sobering. The
Corps of Engineers has come under criticism
for its dredging activities along our coasts
and navigable rivers. Congress authorized a
five-year thorough study of this program that
has now cost $30,000,000. A part of that
study is an eight- volume (1,502 pages) set of
reports covering colonial bird use and plant
succession on dredged material islands. Con-
tracted to seven different teams of qualified
ornithologists, these studies found that "62
percent of all colonial species (more than
156,000 adult birds) along the Texas coast in
1977 nested on dredged material islands." In-
cluded were the Least Tern, the Gull-billed
Tern, the Roseate Spoonbill, the Reddish
30
Great Basin Naturalist Memoirs
No. 3
Egret, and the Brown Pelican. In Florida,
"approximately 50 percent of the colonial
nesting sea and wading birds nest on dredged
material and many more species use the is-
lands for feeding and roosting."
What I gleaned from these studies was the
exciting possibility of so locating and con-
structing these dredge-spoil areas as to create
superior nesting habitat with minimal pre-
dation and disturbance pressure. But the de-
cision between using the most cost-effective
dredge disposal site and a wildlife-oriented
one carries a price tag. For example, to avoid
an endangered plant (Menzies Wallflower)
the alternate to the most cost-effective dis-
posal site for dredge spoil from Humboldt
Harbor (California) is estimated to cost
$150,000.
Some of the requests from the Office of
Endangered Species for these preconsultation
biological assessments pose enormous com-
mitment of time and money. Take the case of
the Nellis Air Force Range in Nevada. BLM
received the following biological opinion:
A study should be conducted to:
(1) Determine all candidate and proposed threatened or
endangered plant species which occur on the Nellis
Range.
(2) Delineate the exact locations of such populations.
Such a study should be for at least one full collecting
season during an average moisture year and prior to any
activities that might jeopardize the existance of the sub-
ject species.
The above instructions for conducting the
study are botanically sound, by reason of the
fact that seeds of many species lie dormant in
the soils of the arid Southwest for years
awaiting an infrequent rain. Then rapidly the
full plant cycle is completed while moisture
is still available. But how do you foretell an
adequate moisture regime? How do you fit
such an indefinite timetable for survey and
site mapping of arid-land plants into the hard
realistic schedules of construction if it is to be
cost effective and available when needed?
Summary: Under the 1978 version of the
act, preconsultation biological assessments
will be the responsibility of agencies seeking
approval of programs authorized, funded, or
conducted by them. The above examples il-
lustrate the potential for delaying the start of
the project and for adding (sometimes signifi-
cantly) to the overall co.sts. It would seem
that regulations have been imposed to pro-
tect animals and plants against extinction be-
fore there is any very precise knowledge of
the tens of thousands of little-known or in-
conspicuous species of nongame animals, par-
ticularly invertebrates, and even less of plant
species we have not chosen to propagate or
value for their form or floral display.
Withdrawal of Resources
There is no way to avoid the commitment
of natural resource if an endangered species
habitat is to be protected. Some of these re-
sources we can easily share, and others are
not in excess of our economic needs. This is
not to say that resources reserved to endan-
gered species are irretrievably lost— but for
current use they are not available, and this
can seriously impact local industries depen-
dent upon them for ongoing supplies.
For example, the U.S. Forest Service has
presently located some 2,000 nesting colonies
of Red-cockaded Woodpeckers in south-
eastern national forests. It has been deter-
mined that each colony nesting site averages
10 acres. One fourth of the colonies require
an additional 25-acre recruitment area. This
is a total of 70,000 acres in merchantable
timber currently removed. The eventual goal
is to have four such colonies per 1,000 acres,
which would entail setting aside 140
acres/ 1,000 acres in suitable timber. There is
an estimated 6,000,000 acres of pine types in
the Red-cockaded Woodpecker's range on
national forests. If the goal is eventually at-
tained, it will mean that 840,000 acres of
commercial timber is being devoted to the
protection of one single endangered species.
While there is no system-wide manage-
ment plan, several regions of the national for-
ests have adopted the practice of setting
aside against any modification eight acres
about each Bald Eagle nest tree, together
with an additional buffer zone limiting activ-
ities during the nesting season. In Alaska
2,760 Bald Eagle nests have been located and
charted, thus automatically setting aside
some 21,500 acres of merchantable timber.
However, land use plans for national forests
in southeast Alaska call for the reservation of
approximately 50,680 acres of standard oper-
able commercial timberlands along beach
1979
The Endangered Species: A Symposium
31
areas, primarily for the protection of eagle
habitat.
The endangered Kirtland's Warbler nests
in northern Michigan in an early successional
stage following fire. Here Jack Pine boughs
screen a ground nest in a more or less con-
tiguous low blueberry ground cover. The re-
covery plan calls for managing some 61,485
acres of Jack Pine on the Huron National
Forest, by controlled burning, to provide this
habitat. Elsewhere, on the Six Rivers Nation-
al Forest in California a proposed timber sale
of approximately 9.25 MMBF of merchan-
table timber was withdrawn to protect an en-
dangered plant (Pine-foot). In New Mexico,
the endangered Jemes Mountain Salamander
requires deep shade and substantial amounts
of moist, decomposing timber material on
rocky north slopes. The management plan
may withdraw as much as 2,500 acres to pro-
tect this habitat, though admittedly the tim-
ber is difficult to harvest. In Montana, habi-
tat protection for the grizzly bear tends to
limit the salvage of beetle-killed timber.
In a number of cases, one of the reasons
given for listing an animal or plant as endan-
gered is overgrazing. However, thus far only
one proposal to close an area to grazing has
surfaced. This is the Beaver Dam Slope area
in southeastern Nevada, for the purpose of
protecting the Desert Tortoise. But with-
drawal of public range can take a number of
forms. For example, prairie dog colonies on
the Buffalo Gap National Grasslands in South
Dakota have increased from 114 in 1968 to
479 in 1975— and the area occupied from
3,000 to 18,000 acres. Because of the endan-
gered Black Footed Ferret that uses the
prairie dog as a principal prey, the simple so-
lution of removing these rodents to the point
where range forage conditions improve is not
acceptable. So the management plan calls for
partial reduction in prairie dog numbers, ac-
companied by a reduction in livestock graz-
ing that would have produced 319,000
povmds of beef.
But of resources in the western United
States that are less than adequate for man's
needs, water stands first. The most produc-
tive place to look for an endangered or
threatened species is in an isolated spring or
sink. Isolation created the adapting species
and that same restricted habitat endangers
them. These sites are very susceptible to
withdrawal of water from underground aqui-
fers for domestic use or irrigation. Thus, the
Desert Pupfish prevailed in stopping a ran-
cher from irrigating his alfalfa fields. In
southwestern Texas three small fish in-
habiting springs and headwaters of drainages
to the Amistad Reservoir are proposed as en-
dangered and/ or threatened, the major rea-
son being "excessive removal of ground wa-
ter." Water uses in an area starved for that
commodity can be affected many miles dis-
tant.
Even cities are vulnerable to this type of
resource withdrawal. For example, to insure
adequate water for future needs, the city of
Cheyenne, Wyoming, acquired the water
rights from the Little Snake River on the
western slope, which they would bring
through a tunnel under the Divide to
Cheyenne. But below the water takeout
points is the stream habitat of the Colorado
Cutthroat Trout, considered for protective
listing. To solve the impasse, Cheyenne
agreed to release 5,000 acre feet of their an-
ticipated 23,000 acre feet of water to main-
tain the trout habitat. The value of the water
to the city is much greater than the $110 per
acre foot necessary to develop the water col-
lection project ($550,000 for this fraction).
The life's blood of the southwestern United
States is the Colorado River drainage basin.
It holds the key to every activity. Endan-
gered species of fish have now been listed for
different segments of this river system from
Wyoming to Arizona. The impact of this pro-
gram in so sensitive an area can be explosive.
Costly Project Modifications
The regional office of the U.S. Forest Ser-
vice in California informed me in August
1978 that they had made 22 requests of the
Endangered Species Office for formal Section
7 Consultations. At that time they had re-
ceived 10 completed biological opinions, half
of which recommended modification of a
program. Similarly, the regional office of the
Forest Service in Montana had received final
biological opinions on five programs, 80 per-
cent of this number recommending changes.
Many of the project modifications were the
product of interagency planning that min-
32
Great Basin Naturalist Memoirs
No. 3
imized cost and disruption. But others add
appreciably to project costs.
The Florida Power and Light Company,
serving the electric needs of southeastern
Florida, is literally being painted into a cor-
ner by a maze of conservation set-aside areas,
including critical habitats for four endan-
gered species. They sought permission to
build a transmission line to cross about a mile
of one comer of the Loxahatchee Wildlife
Refuge. They offered to purchase another
tract of land of equal value that would be
suitable habitat and, in addition, provide $1
million for its development. They failed to
get the easement because it was "in-
compatible with the Everglade Kite Critical
Habitat." The line has now been detoured
around that comer of the refuge at an addi-
tional cost for construction of $1,200,000.
The public utility contends that the easement
they sought contained neither Everglade
Kites nor the Apple Snail on which they feed.
Clear across the continent another public
utility, Southern California Edison, expe-
rienced increased project costs of a somewhat
different nature. A 17-mile equipment haul
route to the San Onofre Nuclear Generating
Station near San Clemente, California, from
the off-loading dock was required. Due to
terrain, land ownership, and load weight con-
straints, the route was to follow along the
coastal beach just above the high tide line.
During 1976, a portion of the route became
populated with a colony of Least Terns. Af-
ter several meetings with the Least Tern Re-
covery Team, it became obvious that a new
haul route and/ or construction schedules and
equipment delivery times would have to be
changed. The studies and altered schedules to
avoid equipment arrival during nesting peri-
od (April-September) resulted in direct costs
of approximately $800,000.
In northern Colorado, the Peabody Coal
Company was enlarging its mining operation,
which is to serve as the sole fuel source for
Colorado-Ute's Power Plant at Hayden.
Peabody had surveyed and purchased a right-
of-way for a haul road to deliver the coal
when a local staff member of the Colorado
Division of Wildlife called attention to a cul-
tivated wheat field along the route that was
used each spring by a small group of Greater
Sandhill Cranes. These migratory stop-over
sites are termed "dancing grounds" because
certain prebreeding rituals take place in this
period. Peabody had prepared an Environ-
mental Impact Study on their program and
circulated it to state agencies, but it evi-
dently did not come to the attention of any-
one knowledgeable about the cranes. The
greater Sandhill Crane is on the Colorado
state list as endangered, but not on the feder-
al. This situation required Peabody to reroute
their delivery road and purchase a new right-
of-way.
The Arkansas State Highway Department,
although filing a formal request for a Section
7 Consultation on the proposed routing of a
four-lane highway, decided independently on
an alternate route to avoid the cave halaitat
of the federally endangered Gray Bat and a
state-listed cavefish and grotto salamander.
The envisioned adverse affects were not the
physical disruption of the right-of-way, but
the off-chance that a chemical spill would
occur on the completed highway that would
enter the undergroimd aquifer that feeds a
more distant cave. This alternate action
lengthened the highway by a little over two
miles, which will cost taxpayers an estimated
$3,000,000.
Addressing Problems, Not Solutions
The Soil Conservation Service has had
some rather difficult experiences with the en-
dangered species legislation. A small water-
shed program has broad participation of af-
fected parties in project planning. The usual
goal is to prevent the loss of topsoil in the
upper basin and destructive flooding in the
lower basin, and to improve permanent wa-
ter sources, be it stream flow or small reser-
voir.
The Cypress Creek Watershed in Lauder-
dale County, Alabama, and Wayne Coimty,
Tennessee, was just such a project. But the
biological assessment that SCS funded turned
up an endangered small fish, the Slackwater
Darter, one of 80 species and subspecies of
darters in Tennessee. The biological opinion
from the Office of Endangered Species point-
ed out that the project would adversely affect
the Slackwater Darter because of its very un-
usual reproductive requirements. While typi-
cal of a slow-flowing stream with silt and
1979
The Endangered Species: A Symposium
33
gravel bottom, this Darter makes use of high
(flood) water to swim off-stream into seepage
areas in open pastures for breeding. OES ap-
proved four water retarding structures on
Little Cvpress where no darters were found,
but blue-penciled for the time being 15 struc-
t\ires on other branches of the system. This
darter needs flooding, but the fields and small
towns down valley don't.
In Mississippi, after identification of the
Bayou Darter in the Bayou Pierre Water-
shed, over $100,000 was spent by SCS to
identify the habitat and critical elements of
that habitat. Planning and taking into ac-
count habitat location and the critical ele-
ments of the habitat resulted in selecting land
treatment and 13 floodwater retarding struc-
tures as the proper approach. An analysis of
impacts on the scope and extent of habitat
and the critical factors in the habitat in-
dicated no significant impact on the Bayou
Darter. But the Office of Endangered Species
disagreed. High on their list of reasoning was
the inability to predict induced land changes
that might be accompanied by increased pes-
ticide residues, siltation, etc. This would not
appear to be an objection to the project per
se, but to the opportunity it provided indi-
viduals within the improved watershed to up-
grade their economic pursuits which just
might adversely affect the darter.
Conclusion
The impacts of the Endangered Species
Act have so many facets and ramify into so
many comers of our society that it has been
impossible in a short paper to bring you very
much of the information finding its way
across my desk in the last three months.
However, it should be abundantly clear that
much of the burden of performing research
and adjusting to endangered species require-
ments is falling outside the coterie of govern-
ment agencies, private organizations, and in-
dividuals who are expressly committed to the
management of wildlife and native plants.
Imposing that obligation places a critical re-
sponsibility on those wielding the legislation
to fully determine that the programs are bio-
logically sound and economically practical.
As the list of endangered species grows, it
will take the wisdom of Solomon to avoid
fencing in the economy until it will no longer
serve you. You have very little time to estab-
lish a favorable rapport, for the program
comes up for another congressional review in
one and one-half years. You have this in your
favor: there is almost no one who doesn't en-
joy some aspect of the living world about us.
Questions to Dr. Spencer
Q. If I interpret your comments correctly and place
them into a context of the relationship they might
have to those of Dr. Clement, there is a real, imme-
diate requirement for changing some of the cultural
practices we presently have. Is this interpretation
correct?
A. I am sorry folks. I live in a pretty practical world
and am not prepared to go into the theories and phi-
losophies of management. So if I may, I am going to
duck that question.
Earlier this morning one of our speakers said that
he was often asked, "What is the value of a given
endangered species and how do you compare it with
the costs that we are going to face in providing pri-
ority-use habitats?" The House/Senate Conference
Report (No. 95-1804, dated 15 October 1978) has this
to say:
... to balance the benefits associated with the agency action against
the benefits associated with alternative courses of action, they should
not balance the benefits of the action against the value associated with
the listed species.
I take this to mean that there is to be no attempt to
place a monetary value on a species threatened with
extinction. In other words, the instruction is to com-
pare the economic impact of the different alterna-
tive actions, but not to place a value on wildlife for
the purposes of comparative costs.
Q. That is correct. It is an act of Congress, I think.
A. It is in the Endangered Species Act Amendments of
1978. The Solicitor General's Office will provide de-
cisions on these matters.
Q. Several of your comments were directed toward the
relative costs of changing a project or altering a proj-
ect in order to be in accord with the Endangered
Species Act. You seem to be saying by this that it
costs a lot of money for other government agencies,
private companies, and the like to accommodate
their designs with the requirements of the Endan-
gered Species Act. I won't argue with that. It's true.
It seems that we need a priority system to go along
with it. As an example, let me tell you a little story.
I had to do an environmental impact statement for a
power line. The question in my mind was, "Is this
power line needed?" I never got a satisfactory an
swer from the power company or anyone else that it
was necessary. It seems to me that we need a prior-
ity system whereby we can feed that kind of infor-
mation into the decision-making process because it
is possible that someone might plan something with-
out a real need for it.
A. I would suggest that this is not a normal procedure.
People generally do not build what they do not
need. It is normal not to encumber an expense un-
34
Great Basin Naturalist Memoirs
No. 3
less you anticipate some beneficial return. Before
being too hasty to ascribe a motive to a person, or-
ganization, or project to which we might have some
objection, I suggest we follow the motto that says,
"Don't criticize your opponent until you have
walked a mile in his moccasins."
The other point I thought you were trying to make
is that people here in the audience ought to be
aware that there is a very bad economic penalty or
economic cost, if you want to put it in those terms,
with this particular piece of legislation and that if
we fail to recognize those costs that are there, we
may be in jeopardy of losing the program entirely.
You have stated my opinion very well. We are proud
of what we have been able to accomplish in wildlife
conservation in this country. Until very recently,
these wildlife programs have been totally self-sup-
porting and have not dipped into the tax till to
which the general public contributes. Now we have
turned around and are progressively passing the
costs along to companies, organizations, and the gen-
eral public for projects in which they have little
first-hand interest. It is up to us to be sure that the
cooperation we ask of them is a wise investment for
all parties.
The new environmental laws, including the En-
dangered Species Act, came into being during a pe-
riod when we were economically well off. Now we
are experiencing a period of inflation, high taxes,
and a cost of living that is affecting every pock-
etbook. It is time for us to be very, very careful we
don't crowd this unfavorable economy. If we ask for
too much, if we wield this powerful legislation with
too much enthusiastic abandon, we stand to have
Congress remove it from the books. Please remem-
ber it comes up in Congress for reappropriation in
18 months. In the 39 years I worked for the federal
government, 34 of those years with the Fish and
Wildlife Service, every time the economy dipped
our appropriations were among the first to be cut. I
don't think times have changed.
There are relatively few endangered and threat-
ened species on the lists at the present time com-
pared to literally thousands that only await the
proper study to be added. We have established criti-
cal habitat for only 33— a not too complicated pro-
cedure when only one species in an area is consid-
ered. But in the future, you can anticipate that
critical habitats will involve acreages and overlaps
that will noticeably fence in the economy.
In my opinion, the 25 amendments to the Endan-
gered Species Act in 1978 succeeded only in making
the legislation more difficult to administer, and
equally more difficult to comply with. It is now so
complex that it is self-defeating.
ENDANGERED ANIMALS IN UTAH AND ADJACENT AREAS
Douglas Day'
.Abstract.— This paper presents a brief background on Utah's experience with the Endangered Species Act of
1973 to date, the Division of Wildlife Resources' involvement with resident endangered wildlife forms, including the
Utah prairie dog, peregrine falcon, bald eagle, woundfin, Colorado squawfish and humpback chub, and problems
associated with the listing of native fauna. Also discussed is a proposal to vest the division with authority for endan-
gered plants by legislative mandate.
I appreciate that kind introduction— and
it's true, I am a son-in-law of Dr. Clarence
Cottam. I was debating whether to mention
that or not, but it has come up. Let me just
pay tribute to Dr. Cottam, as the personal
relationship I had with him was something
special, and I think that the reason I'm here
today is because of the special interest he
showed in me. I can remember looking for
whooping cranes at the Arkansas Refuge. He
wanted to make sure I got a firsthand impres-
sion of those magnificent birds, and that im-
pression has stayed with me throughout my
life. I remember staying out at night with Dr.
Cottam on the Welder Refuge, trying to call
up the Texas red wolf. His keen interest in
endangered species was inspiring. I'll never
forget it. He has been a great influence in my
life. Also, I know he provided some direction
to BYU's biological endeavors.
Talking about endangered species, I think
I might be one. In the position that I'm in as
director of Utah's Division of Wildlife Re-
sources, I think I have a feeling for these crit-
ters that we're talking about. I'm kind of
caught between two worlds— the political
world and the world that we have worked in
so many years in the biological realm. To
make those worlds see eye to eye is very dif-
ficult. That's sometimes why I think directors
are endangered— because they might get a
little too enthusiastic about the biological
part of it and forget the political part. It's a
tightrope to walk. Sometimes we don't have
the opportunity to say what we really feel.
Someone gets to the public before we do and
says this is what they think, and prudence re-
quires that we wait for a better opportunity.
Sooner or later it seems to come. I think the
time might come, if we keep going in the di-
rection we're going in disregarding environ-
mental concerns, that someone might just
happen to have an idea that the whole world
should be declared critical habitat. If that
happens, I don't think we'll have to worry
about collecting permits.
I'd like to make a couple of comments on a
pending court case. The defendants are the
secretary of the interior, the governor of Col-
orado, the director of the Colorado Division
of Wildlife, the governor of Utah, Utah's di-
rector of the Department of Natural Re-
sources, and I. This lawsuit is over threatened
and endangered species. In that lawsuit it is
mentioned that "The right to develop and
beneficially consume the limited quantity of
water . . . (from the Colorado River) is a ves-
ted property right, the use of which is pro-
tected to the citizens, present and fu-
ture. . . ." Now, I would ask the question-
does wildlife have any vested property
rights? I submit that it hadn't, not until the
Endangered Species Act of 1973. That's very
important— to realize why we need to protect
and hold on to the Endangered Species Act.
In that lawsuit some of the claims are the de-
fendants failed to properly, fairly, equitably,
and impartially enforce the provisions of the
Endangered Species Act. Continuing on, the
lawsuit further states, "The factual basis upon
which the determination was made that the
Colorado Squawfish and the Humpback
'Division of Wildlife Resources, 1596 West North Temple, Salt Lake City, Utah 84116.
35
36
Great Basin Naturalist Memoirs
No. 3
Chub are or were 'threatened with extinc-
tion,' and the continued designation under
ESA as 'endangered' was and is not based
upon sound and adequate biological data and
knowledge of said species . . . and, further,
amount to arbitrary and capricious acts on
the part of the defendants. ..."
Here is another item, "The defendants, and
all of them, have determined without ade-
quate biological data and knowledge that wa-
ter impoundment development adversely af-
fect such fish species. . . , and as a result of
said erroneous conclusions, based upon little
or no scientific evidence, defendants have
continued to wrongfully impede plaintiff dis-
tricts' efforts to construct their projects, in-
cluding impoundments. . . ."
And last, another excerpt I thought would
be of interest to you, is "The fact that Colo-
rado Squawfish, and the Humpback Chub,
were allegedly 'threatened with extinction'
and are now allegedly 'endangered' is the di-
rect and proximate result of the stocking by
all defendants of non-endemic, non-native or
exotic fishes in the Colorado System." What
that means is that the stripers are eating the
squawfish. I suggested a proposition or a pro-
posal that might be humorous in one way but
sad in another, that being, if they really want
to stop the stripers from eating the squawfish
(there is no scientific evidence of this), why
don't they build another dam on the Colo-
rado River to keep stripers from running up-
stream. I don't know what the outcome of all
this will be.
My time is rather limited, but I want to
draw your attention to last Sunday's Parade
magazine. I am pleased that we are getting
this kind of coverage. What it says is the two
things that are the greatest threat to wildlife
today are (1) loss of habitat (and that's very
obvious, because if endangered species had
good habitat they wouldn't be in danger) and
(2) commercialization of world wildlife. Con-
sider these statistics from the U.S. Fish and
Wildlife Service. Last year in the United
States we imported about 100 million tropi-
cal fish, 500,000 reptiles, 100,000 mammals,
and uncounted thousands of birds. The im-
ported traffic in manufactured wildlife
goods— furs, coats, leather, trinkets, jewelry,
and carvings— leaped from 1.7 million items
in 1972 to 91 million in 1976, the last year
for which figures are available. Between
1973 and 1976, skin and hide imports rose
from 900,000 to 32.5 million. Part of the rea-
son is the impact from TV of Barretta's bird,
Fred. A few years ago you could buy a cock-
atoo for $100, and now some of them are
fetching $6,000 apiece.
Let me go into some of our involvement in
the State Wildlife Division with endangered
species. I'll tell you about a few of the spe-
cies we're working with and about some of
the progress we are making. But first, I want
to take just a minute and maybe leave you
with another concept. I've worked closely
with the Boy Scouts for a number of years,
and I had the opportunity to take them to a
power plant. One thing that impressed us
was the control room. In that control room
you could virtually feel the whole operation
of the plant. It was right there; you knew
what was happening, and when there was a
problem somewhere a red light came on. The
plant operator could tell where that problem
was from the red light.
I would submit to you that in the biologi-
cal world we have our red lights. We don't
pay much attention to them, or haven't done
until lately. These red lights are our endan-
gered species. I think that is a good concept.
I noticed yesterday morning driving to the
office a pickup truck that I was following
was obviously losing its antifreeze, and I
could predict what would happen; the red
light came on and the truck was in trouble.
He could go on a little while, but eventually
it had to be taken care of or that truck was
doomed. The operator obviously paid atten-
tion to the warning light and pulled off the
road.
In the biological world we don't pay atten-
tion to our red lights as we should. We're just
beginning to do this. These are our endan-
gered species. This much-used and publicized
terminology connotes a wildlife form desper-
ately trying not to join the passenger pigeon,
heath hen, and others in the land of memory.
Each time we lose a species one of our red
lights goes out.
This designation of endangered species has
also been accused of holding up progress and
projects, locking up land, and various other
alleged abuses. The Endangered Species Acts
of 1966 and 1969 were relatively innocuous
1979
The Endangered Species: A Symposium
37
in that they recognized the status of certain
species and hsted them, but it took the En-
dangered Species Act of 1973 to estabhsh a
national policy to come to grips with the is-
sue and determine ways and means to at-
tempt to reverse the trends of certain species
toward extinction.
To a state wildlife administrator, the 1973
Act with its attendant rule making and re-
strictions has been the source of much soul
searching. I believe most of us whole-
heartedly agreed with the philosophy and the
intent of Congress. We have vigorously ob-
jected to the early federal agency approach
that absolutely usurps state authority in en-
dangered species management. Recent devel-
opments have ameliorated the situation, and
on the horizon I can see finally the devel-
opment of a much closer state-federal work-
ing relationship with the goal of doing all hu-
manly possible to restore endangered species
to a viable component of our environment.
The sadness in the situation is that it has
taken almost five years to get to this point.
This is time that we can't afford to waste.
However, in defense of the federal agency's
past position, let me add that the act itself,
until amended, left no room for legitimate
compromise. This in itself has been a big
stumbling block.
I remember another Cottam that you are
well acquainted with here at BYU, and that is
Dr. Walter Cottam. He said, "Unless you
learn somehow to compete with the dollar,
you will lose the conservation battle." In my
experience, and ever since I have been in-
volved, it has been a compromise situation. I
am afraid the direction we are going is com-
promise to extinction unless we reverse that
trend. We are just beginning to get some
tools that give us a little bit of an edge in the
compromise situation. Because of the devel-
opmental demands in our environment, it is
not easy to carry on this struggle we are in.
Believe me, it is discouraging. I can remem-
ber only one instance when a developer came
on his own to a wildlife biologist for input
into a development project. It is sad that we
have endangered species acts and other legis-
lation to require coordination and con-
sultation between developers and biologists.
The real plus for the act has been the
awakening interest in the amount of knowl-
edge about many species we never before
considered as significant, or for that matter as
ever existing. In retrospect, our formal train-
ing in the field was deficient in many areas
but suited the times. Unfortunately, industri-
alization, social pressure, and human de-
mands accelerated at a rate faster than the
state of the art of wildlife management. Re-
lated fields of plant and animal science have
produced knowledgeable individuals who
have "come out of the woodwork," so to
speak, with indisputable evidence regarding
certain species that state management
agencies were never privy to, were unaware
of, or disregarded.
For many years we have been game orien-
ted, not always by choice, but by the unre-
lenting force of simple economics. Until
1975, in Utah, our entire program was fi-
nanced by user fees in the form of hunting
and fishing licenses, fines and forfeitures,
matching federal monies also paid by hunters
and fishermen, and miscellaneous sources. It
is obvious our primary mission has been to
provide for and produce those species sought
after by those paying the bill. In 1975 the
Utah Legislature provided general funds to
iinplement a modest nongame section within
the division and has continued that support,
still modest in terms of total budget. There is
a legitimate need to increase funding for non-
game programs, to increase our capabilities
to provide basic knowledge and solutions to
current problems. Appropriate emphasis is
being placed on endangered species within
this nongame section.
With this backgroimd, let me detail pro-
grams related to endangered species in Utah.
First, let me say that we have yet to enter
into a formal cooperative agreement with the
U.S. Fish and Wildlife Service under terms of
the Endangered Species Act. Recent congres-
sional action provides for new rule making
allowing us to do this, and we expect to sign
such an agreement. However, the lack of a
formal agreement dampened neither our ded-
ication nor enthusiasm to get on with the job
that needed doing.
In 1973, under a special cooperative agree-
ment with the Denver Regional Office of the
U.S. Fish and Wildlife Service, funds were
provided to survey historical and potential
habitat of the Utah prairie dog {Cynomys
38
Great Basin Naturalist Memoirs
No. 3
parvidens). This is our start. As an endan-
gered species, this animal has had an erratic
history. It was first added to the hst in 1969,
removed in 1970, and added again in June
1973. Since our initial effort in identifying
habitat, the original agreement has been
amended annually, providing funds each year
to continue our trapping and transplanting
programs. In spite of its endangered species
status, based on overall population and status
of colonies, those found in private agricul-
tural lands have provided us an annual prob-
lem of some magnitude. It is from these dam-
age situations that we trap and attempt to
establish new colonies in areas of historical
occupancy on public lands. Trapping com-
mences in the spring before the young are
born, ceases until young are weaned, and re-
sumes and continues until late summer. Our
most recent technique attempts to livetrap
family groups for relocation. The work is car-
ried out using seasonal employees supervised
by our regional office in Cedar City under
guidelines from our nongame section.
We have come under criticism from one
animal protection organization for what ap-
pears to them to be a low success rate of sur-
vival and establishment from our transplant
program. Also, that organization is critical of
our numbers for the species as compared to
the estimated population in 1973. All I can
offer is that, because we are not sure of a
percentage to project for a total population,
we will continue to use our maximum counts
as a minimum population figure. I am sure
that our sincere efforts to alleviate agricul-
tural damage has kept some landowners from
taking matters into their own hands. Even if
we are losing large numbers in an attempt to
establish a new colony, it appears to us to be
a wiser decision than to possibly lose the
same number or more without having taken
the risk. We are not happy with the odds ei-
ther, but restoration of any species is fraught
with failures, disappointments, and frustra-
tions. We are only human in recognizing and
being affected by them but feel that we are
also professional in not giving up and in gen-
uinely trying to reduce these failures, dis-
appointments, and frustrations. We are con-
fident that the Utah prairie dog will soon be
in a secure enough position from the stand-
point of new colonies on public lands that we
can successfully petition for delisting. What
we attempt to do is investigate the problem,
do the research necessary, and give some
management that will ensure an environment
in which the species can live and reproduce
and have some kind of continuance. It takes
management.
In May 1975, before authorized funding
for our nongame program became effective,
we concluded, in an agreement with the U.S.
Fish and Wildlife Service and Bureau of
Land Management, to jointly fimd the divi-
sion position of raptor biologist. Our share
was fimded through monies received as con-
tribution from private citizens. This arrange-
ment continues to this day, except that our
share has been funded by legislative appro-
priation since 1 July 1976.
Our work with raptors includes all species,
with emphasis on those endangered, sensitive,
or unique. The American peregrine falcon
{Falco peregrinus anatum) is of primary con-
cern in view of its current national status.
Based on historic records, Utah had the high-
est rate of occupancy by this subspecies of
any western state with the possible exception
of California. You are all aware of the dra-
matic decline in the West and the extirpation
of the falcon in the East and the possible
causes. From the middle sixties to 1975 there
were no known active peregrine eyries in
Utah; at least none were revealed to us. In
1978, we documented occupancy at four
sights, but fimding limitations allowed no
data to be gathered through our efforts— that
was because of a lack of personnel. We do
what we can. Whether the peregrine is stag-
ing a comeback is a matter of conjecture. We
doubt it. Increased awareness of its plight is
probably the reason for recent docimienta-
tion, plus limited additional effort to seek out
the presence of the species. Evidence avail-
able indicates pesticide residues are still too
high to cause much optimism at this point.
We will continue to put as much effort to de-
termining status as fimds and personnel will
allow. This activity will increase when En-
dangered Species Act funds become available
to us.
This year the bald eagle {Haliaeetus leu-
cocephalus) was added to the list of U.S.
threatened and endangered species. Pre-
viously only those bald eagles nesting south
1979
The Endangered Species: A Symposium
39
of the 40th parallel were listed. This action
has caused us no little concern because we
have no documentation of nesting bald eagles
in Utah, but each winter we are visited by
over 600 eagles produced from as far away as
northern Saskatchewan. The population
seems thriving and secure there, and we are
at a loss to explain how their plight changes
as they wing their way over a political
boundary. I am reminded of a settler who
was living up in that area along the United
States-Canadian boundary at the time they
surveyed our modern boundary lines. He
found out through the survey that he was ac-
tually in the United States. He said, "Thank
God. I couldn't have stood another Canadian
winter. "
We protected the listing for several rea-
sons, but the one of the greatest magnitude
and potential problem is that of critical habi-
tat designation if the action ever comes to
pass. Every canyon used for night roosting,
every tree used for day resting, every hunting
area could come under the designation. What
would happen to our waterfowl management
areas upon which so many nonhunted species
depend if federal funds were withheld for
failure to limit hunting because of the pres-
ence of bald eagles? Today there hasn't been
a hint of such action, but stranger things have
happened. Before the recent listing, we in-
itiated a survey of bald eagle visitants, and
for the past five years have documented
number, preferred location of use, and gener-
al arrival and departure dates. This year, in
addition to our own winter census, we will
participate in the national one-day bald eagle
census in cooperation with the National
Wildlife Federation.
I want to talk for a minute about one other
species, mention some fishes, and then wind
it up. The desert tortoise {Gophenis agassizi)
was mentioned earlier this morning. Recently
the U.S. Fish and Wildlife Service has pro-
posed, by Federal Register publication, listing
of the desert tortoise as an endangered spe-
cies along with designation of 38 square miles
of the Beaver Dam slope in Washington
County as critical habitat. We have been ac-
tively involved in recent years in document-
ing the current status of the tortoise and the
condition of its habitat. While sympathetic to
its plight, we believe this move to be pre-
mature in that studies currently under way in
Arizona immediately adjacent should be fin-
ished and evaluated and the entire system
looked at rather than drawing political
boundaries to attempt management of a spe-
cies.
I will touch only briefly on the endangered
fishes here in Utah. I am sure Dr. Deacon
will provide more in-depth summary in his
presentation. There are presently three spe-
cies in Utah in this category. They are the
Colorado squawfish {Ptychocheilus lucius)
and humpback chub {Gila cijpha) in the
mainstream Colorado and Green rivers in
Utah and the woundfin {Plagoptenis argentis-
simus) found in the Virgin River below La-
Verkin Springs. There are three more species
currently under consideration for either
threatened or endangered status— the razor-
back sucker {Xyrauchen texanus) and bony-
tail chub {Gila elegans) in the mainstream
Colorado system and the Virgin River round-
tail chub {Gila robusta seminuda). Our past
work with these has been very limited, pa-
ricularly with the Virgin River species. Re-
cently, we have been more involved and ex-
pect to fulfill our role as fish and wildlife
managers as funds are made available.
It now appears our next step may be into
the area of endangered plants. In a few min-
utes you will hear more of the status of cer-
tain plant species from persons more knowl-
edgeable than I; however, the Utah Science
Advisory Committee has prepared legislation
for introduction at the general session of the
legislature in January that will give our divi-
sion jurisdiction over those plant species de-
clared threatened or endangered under the
act. It also provides for the establishment of a
position of taxonomist and funding to carry
out the necessary activities. If this comes to
pass, we will be drawing heavily for some
time on the expertise of several of the speak-
ers at this symposium.
Our involvement with endangered species
to this point, though not deep, has been sub-
stantial considering the minimal funding re-
ceived imder the act for just one facet of the
program. Recovery teams are in operation
for all Utah species, and we have representa-
tives for all but the bald eagle. Our participa-
tion has been active and sincere. We take the
task seriously and intend to continue to pro-
40
Great Basin Naturalist Memoirs
No. 3
vide meaningful input for the sake of the spe-
cies involved. We also intend to cooperate
with other states, federal agencies, and all in-
terests to ensure that endangered species are
provided for, keeping in perspective the
needs of all wildlife as well as those various
interests of our human resource. I hope our
efforts will be interpreted in this light.
Questions to Mr. Day
Q. We agree with your present analysis of the situation.
If you find a better solution, please let us know.
A. I think what we in wildlife management have to do
is to make intelligent decisions concerning endan-
gered species and keep the pendulum from swinging
too far the other way. We don't want to lose this
tool, as I mentioned before, and I think you can see
what effect the politician has on the direction we go.
Q. I'm not sure where these big birds move in from, but
in the Uinta Basin there's quite a wintering popu-
lation of bald eagles which comes in and feeds on
the waterfowl of the Pacific flyway that goes
through that area. There is an area there that came
to be recognized as a roost area where the birds go
back and rest in the evening and spend the night.
Such areas have been given refuge status in Oregon.
A. Well those things happen. We need to use prudence
in recognizing these areas or things can get out of
hand.
Q. Do I sense an opposition to listing any critical habi-
tat in Utah?
A. From our division? We don't list it. We are not op-
posed to critical habitat designations if needed.
Q. Well, Dr. Murphy said the same thing there and I
was wondering if . . .
A. No, when you get down to specific cases, whatever
is needed, whatever the facts require to ensure the
.survival of that species at an intelligent level— that
ought to be the way we are managing it. You can see
what the reaction of the public and the politician is
to situations where we go overboard.
Dr. Murphy: There are several large and impor-
tant roosts that would fit the category of the one he
mentions in the Uinta Basin that I would be very
happy to see listed as critical habitat, but it's just
that the wintering population becomes very dis-
bursed and small groups will be found in small areas
all over the state. It becomes almost administratively
impossible to keep up with that kind of a situation.
Q. I would like to ask you a question that is perhaps out
of your realm, and that is "what is the policy of the
state with regard to endangered species, particularly
plants on state land?"
A. A lot of people have been asking me those kinds of
questions lately. I guess first we'd have to know
what the management implications are. You've got
the other species I mentioned, the resources, the re-
habilitation projects going on for game species, and
that type of thing. I can only answer, just in a gener-
al way, that we're interested. We want to see these
identified and take intelligent mea.sures to protect
habitat and species.
Q. Specifically, what about the Cactus rideii on the
Mancos Shale in the Citros Butte area of Wayne
County which is being strip mined for coal and is in
an area of critical habitat for that species?
A. You remind me of a story that will maybe get me off
the hook. This fellow was a well-known speaker. He
gave this talk, and his chauffeur drove him around to
all these places, and his chauffeur said one day, "Let
me give this talk for you. I've heard it so many times
I can do it as well as you can."
So he said, "OK, I'll wear your chauffeur's uni-
form and you give the talk."
That happened. The audience applauded, and
then it came time for the questions. That's the situa-
tion I'm in. A question like this came up, and he
said, "That's one of the simplest questions I've ever
heard, and to show you how simple it is, I'm going
to let my chauffeur answer it." The problem is that I
didn't bring my chauffeur.
We'll work with you. Let me just indicate that
state lands are not our wildlife lands, but lands un-
der the State Land Board. I'm not happy with the
past management of state lands. Overgrazing has
been a continual problem since early in Utah his-
tory. We're stuck with the rehabilitation. I've seen
that overgrazing. The most recent time was the bow
himt this fall on the Manti Forest. Y'ou can look at
the museum pictures of overgrazing and you can go
out on Fred's Flat today and identify those same pic-
tures without a camera. If you don't learn from his-
tory, you're bound to make the same mistakes.
ENDANGERED AND THREATENED FISHES OF THE WEST
James E. Deacon'
,\bstract.— The endangered and threatened fish fauna of the United States exhibits problems resulting primarily
from habitat modification by man. The evolutionary history of the fauna has left it especially sensitive to biotic
interactions. In addition, many forms are of such restricted distribution that the entire taxon can be destroyed by
very minor perturbations. The effects of habitat modification on woundfin and roundtai! chub in the Virgin River of
Utah, Arizona, and Nevada are discussed. Parasitism by Lernea on White River springfish is shown to coincide with
population decline in some, but not all, cases. Population declines of Pahnunp killifish are related to biotic inter-
actions with both goldfish and mosquitofish. Population size of Devils Hole pupfish are shown to be quite responsive
to small changes in habitat availability.
Fishes of the West are affected by the same general kinds of ecological problems that are causing extinctions
throughout the world. The interplay of economics with perceived value in society has led us into the numerous
ecological problems facing us today. There is some evidence to suggest that society is making some preliminary ef-
fort to slow the rate of extermination. Perhaps this is happening because the conclusions of ecologists, philosophers,
and theologians regarding the relationship of man and environment are to sopie extent being translated into legisla-
tion as well as into conventional wisdom.
The fish fauna of the western United States
has frequently been characterized as one hav-
ing a relatively low diversity and containing
an unusually high percentage of endemic
taxa exhibiting limited distributions (Miller
1959, Smith 1978). These appear also to be
the primary features contributing to the fact
that much of the fauna is threatened to some
degree.
Recently, the Endangered Species Com-
mittee of the American Fisheries Society
compiled a listing of threatened fishes of
North America (Deacon et al. 1979). The
fishes on that list from the western United
States are presented here as a data base for
the general discussion (Tables 5 and 6). The
predominant threats to all taxa listed were
generalized into five broad categories and
each taxon was assigned one or more of these
categories. Threat categories were as follows:
(1) The present or threatened destruction,
modification, or curtailment of the habitat or
range. (2) Ovenitilization for commercial,
sporting, .scientific, or educational purposes.
(3) Disease or parasitism. (4) Other natural or
manmade factors affecting continued exist-
ence (hybridization, introduction of exotic or
translocated species, predation, competition).
(5) Restricted range of the taxon. A com-
parison of threats to western fishes north of
Mexico with those to eastern fishes is of gen-
eral interest and illustrates significant differ-
ences between the two faunas (Table 1).
Habitat modification (Category 1) is clear-
ly the most prevalent threat to native fishes
throughout the world, and this is certainly
tRie in North America. There are a few spe-
cies in the West, however, that are not now
so threatened. No eastern species, however,
has escaped problems raised by physical al-
teration of the habitat.
No western species has been or is threat-
ened by overexploitation (Category 2), but
about 7 percent of the eastern fishes on the
list are or were so threatened. Six species of
ciscoes occurring in the Great Lakes were
subjected to overfishing by commercial fish-
ermen, changes resulting from the in-
troduction of the sea lamprey, and general
environmental degradation (Scott and Cross-
man 1973). In addition, the Atlantic whitefish
has been subjected to overfishing as well as
habitat alteration. They represent the only
fish taxa in the United States or Canada to be
on the American Fisheries Society list of
threatened species, in part, because of over-
exploitation.
Di.sease and parasiti.sm (Category 3) have
'Department of Biological Sciences, University of Nevada, Las Vegas, Nevada 89154.
41
42
Great Basin Naturalist Memoirs
No. 3
apparently not been involved in threats to
any eastern species on the list but have been
factors for about 4 percent of the western
fishes. It is probable that this difference re-
sults from the fact that information regarding
incidence of disease and parasitism in native
fishes is relatively sparse. In addition, though
the initial major decline in abundance and
distribution of eastern fishes probably oc-
curred prior to 1850 (Trautman 1957), in the
West the similar event occurred subsequent
to 1850 (Miller 1961). Because increased in-
cidence of disease and/ or parasitism as an
important factor in a population decline be-
comes most apparent during the major de-
cline, it must be detected at that time to be
recognized. The generally earlier decline of
eastern fishes during a time when increased
incidence of disease or parasitism would have
been less likely to have either been detected
or associated with the decline probably ex-
plains its absence from association with the
eastern faima. This factor doubtless has been
a more important contributor to decline of
both eastern and western fish populations
than is apparent. It has specifically been
identified by Wilson et al. (1966) and Seetha-
ler (1978) as a factor in the decline of west-
em fishes.
Biological interactions of various kinds
(Category 4) contribute to the problems
faced by 54 percent of the threatened west-
ern fauna but only 9 percent of the threat-
ened eastern fauna. The marked differences
in Category 4 point to distinctions of the
western fish faima that have been repeatedly
discussed. Physical barriers to dispersal have
resulted in relatively low colonization rates
throughout the West, with the consequence
that western fish faunas are not especially
speciose (Smith 1978). Because their evolu-
tionary experiences have been with relatively
depauperate faimas, western fishes have rela-
tively low tolerances to biological inter-
actions (Smith 1978, Deacon and Minckley
1974, Hubbs et al. 1974).
A restricted range (occurring in only a
single spring, a single group of springs, or a
short stretch of stream [Category 5]) is a fac-
tor involved in giving a threatened status to
21 percent of the western fishes listed, but
only about 7 percent of the eastern fishes.
Category 5 illustrates the fact that one group
of western fishes appears to have a high de-
gree of "extinction resistance" (Smith 1978).
The consequence is that many western taxa
exist as relict populations in single habitats.
They found their way onto the AFS list of
threatened fishes because of that fact. They,
like many western fishes, generally have high
tolerances to physical extremes but low toler-
ances to biological interactions (Deacon and
Minckley 1974).
Physical Modification of Habitats
While western fishes have in general de-
veloped considerable resistance to the phys-
ical extremes imposed upon them by climatic
factors, they have also been most strongly af-
fected by general and specific alterations of
physical habitats imposed upon them by
man. Miller (1961), Hastings and Turner
(1965), and Cottam (1961) have dramatically
shown the impact of slight climatic shifts su-
perimposed on removal of vegetative cover
by overgrazing between about 1880 and
1900. The arroyo cutting, siltation, and de-
watering that occurred during this period
were probably the most detrimental 20 years
Table 1. Comparison of general kinds of threats to the threatened freshwater fish fauna of western and eastern
North America, north of Mexico.
General threat category
Western Fishes
Eastern 1
s-ishes
Number
Percent
Number
Percent
of taxa
of fauna
of taxa
of fauna
(N = 112)
affected
{N = 90)
affected
109
97.3
90
100
0
0
6
6.7
5
4.4
0
0
60
54
8
8.9
24
21
6
6.7
1. Habitat modification
2. Overexploitation
3. Parasitism and disease
4. Biotic interactions
5. Restricted range
1979
The Endangered Species: A Symposium
43
of all time to fishes and aquatic habitats in
the western United States. This period was
followed closely by a very active period of
dam building, with concomitant increases in
irrigated agriculture, especially since about
1930, when large reclamation projects began
providing water to irrigate what is now some
10 million acres of land in the West. The de-
cline in abundance of the native fishes of the
mainstream Colorado River is associated
closely with construction of these mainstream
dams (Minckley and Deacon 1968, Holden
and Stalnaker 1975 a, b, Seethaler 1978). De-
clines in fishes of tributary streams are also
occurring and are similarly associated with
water manipulations of various kinds that re-
sult in dewatering portions of fish habitats.
Recently, McNatt (1978) has described the
process along the San Pedro River of Ari-
zona. I present some documentation here for
similar problems along the Virgin River of
Utah, Arizona, and Nevada.
The Virgin River drains southwestern Utah
and flows through the northwestern comer of
Arizona before joining the Colorado River in
Lake Mead, Nevada. A salt spring, LaVerkin
Springs, enters the river 180 km upstream
from its confluence with Lake Mead, forming
the upstream limit of distribution for both
the Virgin River roundtail chub, Gila robusta
seminuda, and the woundfin, Plagopterus ar-
gentissimus. Both are here listed as endan-
gered and both are presently restricted to the
mainstream of the Virgin River below LaVer-
kin Spring. In addition, the Virgin spinedace,
a threatened species, occurs both below and
above the springs.
Irrigation diversions have been established
along the river since the 1860s. Since at least
the early 1900s, the Hurricane Diversion,
Washington Diversion, and Mesquite Diver-
sions (Fig. 1) have been capable of diverting
essentially the total summer flow of the river
at each of these three diversion points. La-
Santa Clara River
Boulder Dam
Wash
Mesquite
Diversions
Washington Diversion
UTAH
LaVerkin Creek
Hurricane Diversion
Lake Mead
Fig. 1. Mainstream Virgin River below Hurricane diversion showing total remaining potential habitat for the en-
dangered woundfin and roundtail chub, and significant modifications currently restricting their range.
44
Great Basin Naturalist Memoirs
No. 3
Verkin Springs, entering just below the Hur-
ricane Diversion, plus inflow from LaVerkin
and Ash Creeks, maintain permanent stream
flow downstream to Washington Diversion
(Fig. 1). Littlefield Springs, entering at the
lower end of the narrows, maintain per-
manent streamflow downstream to the Mes-
quite Diversion (Fig. 1). When the total
streamflow is actually used at the above di-
version points, only about 52.5 km (or 29 per-
cent) of the remaining 180 km of potential
habitat for the two endangered species re-
stricted to the mainstream is actually con-
stantly available to them.
The narrows (Fig. 1) divides the main-
stream into an upper and a lower component
that appears to effectively isolate the con-
tained fish populations. Elevation and cli-
mate in the two regions differ significantly.
The difference was reflected by the nearly
one-month earlier spawning of the woundfin
population in the lower river in the spring of
1977 (Fig. 7).
The question of requirements of these fish-
es in their remaining habitats has been the
subject of studies conducted at various levels
of intensity since 1961 (Cross 1975, 78, Wil-
liams 1977, Schumann 1978, Peters 1970,
Lockhart 1979, Vaughn Hansen Associates
1977). The drought of 1977 resulted in some
of the lowest flows on record in the Virgin
River, a circumstance which allowed signifi-
cant insights into the probable effects of wa-
ter development projects which would tend
to reduce or alter flows in the river. The
more normal flows of 1978 provided a useful
comparison to the low-flow conditions of
1977.
Length-frequency analysis was used as a
convenient means of examining the popu-
lation structure of the fishes in the Virgin
River. Samples were taken by repetitively
seining an area until the number of fish col-
lected amoimted to less than 10 percent of
the highest number collected. In this way we
insured a good representative sample of all
fish occurring in the sampled area. Figure 2
demonstrates that samples taken in August
1977 and more extensive sampling from No-
vember 1977 provide essentially the same
picture of population structure for woundfin.
This suggests that sampling done in both Au-
30.
25.
20.
15-
10.
23Aug. 1977 (N= 70)
25-26 Nov. 1977 (N= 370)
18 20
/\ry^^
30
40
50
90 SIZE
Fig. 2. Length frequency of woundfin in Virgin River above the narrows during fall 197
1979
The Endangered Species: A Symposium
45
gust and November was extensive enough to
provide a good representation of population
structure in woundfin. The major fact re-
vealed is that in 1977 young-of-the-year com-
prised a very small (nearly inconsequential)
proportion of the woundfin population above
the narrows. By contrast, a comparison of
population structure in woundfin above the
narrows in 1977 and 1978 (Fig. 3) indicates
that young-of-the-year dominated the popu-
lation in 1978.
When sampling is extensive enough, and
stunting can be discoimted as a significant
factor, much of the information gleaned from
an examination of length frequency can be
summarized by calculation of a mean length
for the population. In this case, for both
woundfin and roundtail chub, small mean
length indicates relatively high reproductive
success and vice versa. Figure 4 and Table 2
present data available on mean length of
woundfin above the narrows in 1973, 1977,
and 1978, together with a hydrograph of
mean monthly flows. They show that in 1973
and 1978, with high winter and spring flows,
reproductive success was high, but in 1977,
with low flows, reproductive success was
low.
A similar situation appears to have existed
for the roundtail chub, Gila robusta semi-
niida (Fig. 5, Table 2), except that the species
was so rare in 1977 that very few were cap-
tured in spite of extensive sampling efforts.
This, of course, indicates that not only were
environmental conditions in Virgin River
during 1977 inimical to successful spawning
in this species, they also apparently reduced
the survival of adults. Figure 5 does show
that the species spawned successfully in at
least one location on the upper mainstream
of the Virgin River in 1978. Relatively high
population density or evidence of a successful
hatch was not found at any other location
sampled in the upper or lower Virgin River
30.
O-O FALL 1977 (N=370)
^-* FALL 1978 (N = I53)
25-
POPULATION
1
o
/
^ 10.
/
5.
/
^
<^x^/
,><T , -Ji , ^ *,
=^:=-i?
20 30
40 50 60 70 80 90
SIZE
Fig. 3. Comparison of length frequency of woundfin in Virgin River above the narrows in fall 1977 and fall 1978.
46
Great Basin Naturalist Memoirs
No. 3
Table 2. Mean size of woundfiii and roundtail chub in Virgin River. The 0 indicates collections were made in the
area but no individuals of the species were taken. The — indicates the area was not collected. Data on woundfin
from 1973 were provided by Mr. Jerry Lockhart. He probably also took chubs; however, data are not available.
1973
1977
1978
Aug &
Sept
1-8
June
2.3-30
Aug
14-15
Nov
25-26
Nov
12
April
28
Sept
1
Nov
X
N
X N X N X N X N
X N X
N X N
Upper river
Woundfin
52.1
60
69.3 172 74.3 70 71.2 416 72.3 .371
76.2 190 46.0
153
Chub
158.2 5 173 1 0 1
0 67.6
102
Lower river
Woundfin
50.2
105
44.0 177 53.6 202 - 56.6.383
58.2 112 44.4
427 48.6 46
Chub
45.0 1 173 1 - 158 3
0 84.3
4 70.0 1
in September 1978. Perhaps even in times of
"normal" flows there are relatively few op-
timal habitats for the roundtail chub remain-
ing in the Virgin River.
In addition to the marked differences in re-
productive success of woundfin and roundtail
chubs in 1977 and 1978, interesting differ-
ences in population structures of woundfin
above and below the narrows in 1977 were
evident. It is apparent from an examination
of Figure 6 and Table 2 that young woundfin
comprised a far greater proportion of the
woundfin population in the lower river in
1977 than was true in the upper river. The
effect of the drought on woundfin population
structure in the upper river thus appears to
have been more severe than was true in the
lower river. In the lower river, however, it is
apparent that by fall and winter 1977 older
or larger fish tended to predominate to a
greater extent than was true in either fall
1973 or fall and winter 1978. This suggests
(1) that growth of young in 1977 may have
been faster than was the case in 1973 and
1978, (2) survivorship may have differentially
favored older woundfin during summer 1977,
(3) spawning may have occurred earlier in
summer 1977 than in 1973 or 1978, or (4)
perhaps more probable, the later secondary
spawning period was almost entirely unsuc-
cessful in 1977, whereas in 1973 and 1978 it
was successful and resulted in a significant
contribution to the population in the fall
(Fig. 1). The effect of the two spawning peri-
ods in 1978 is apparent in Figure 2 as two
peaks in the length-frequency diagram in the
size range below 60 mm.
Figure 8 shows dates on first appearance of
10.000
1000
-3; 100
90
80 3
CD
O
70 =>
I- 60"*
50
[ 40
1973 1977 1978
Fig. 4. Mean monthly flow at Hurricane Gage and mean size of woundfin in upper Virgin River 1973, 1977, 1978.
1979
The Endangered Species: A Symposium
47
woundfin fry at various locations along the
Virgin River during summer 1977. Collec-
tions were made at one- to two-week inter-
vals until fry were taken at each location in-
dicated. The uppermost location, indicated
by 23 July in Figure 8 was actually below the
Hurricane Diversion but above LaVerkin
Creek. It is apparent that hatching occurred
earlier in the lower river than in the upper
river. Furthermore, in the lower river hatch-
ing appears to have been delayed by about
two weeks at the lowermost station where
habitat modification is most obvious.
The earlier appearance of young woundfin
in the Arizona segment of the lower river
was followed by relatively good survival in
1977 (Fig. 7). By contrast, the later appear-
ance of young woundfin in the upper river
was followed by very poor survival in 1977
(Fig. 3). With higher' flows in 1978, both up-
stream and downstream populations of
woundfin showed good reproduction, and by
fall 1978 the mean size was nearly identical
in the two populations (Fig. 6).
Comparisons of hydrographs of Virgin Riv-
er flows for 1973, 1977, and 1978 show that
the major differences in flow occurred during
winter and spring. Summer flows suggest a
relatively greater degree of similarity for all
three years (Vaughn Hansen Associates 1977).
If winter and spring flows significantly in-
fluence reproductive success of the endan-
gered fishes of the Virgin River, the effect
should be discernable in the population struc-
ture during the following fall. Figure 9 pres-
ents data comparing mean size of woundfin
in the fall in both the upstream and down-
stream populations against mean flows of the
river during the spring. Of particular signifi-
cance is the fact that when stream flow is
low, mean size is high and vice versa. Inter-
estingly, Figure 9 also suggests that when
mean spring flows are above 700 cfs, repro-
ductive success may be slightly poorer than
when mean spring flows are between 400 and
600 cfs. Data are not available for times
when mean spring flows fall between 100
and 400 cfs, but at about 100 cfs it is clear
25-
9/28/78 (N=I02)
11/25/77 (N= 3)
2^_^
— I—
150
— r-
180
210
240
270
300
SIZE
Fig. 5. Length frequency of roundtail chub in Virgin River above the narrows. The o's indicate size of the only
three individuals taken in extensive sampling on 25 November 1977.
48
Great Basin Naturalist Memoirs
No. 3
that reproductive success falls off dramatical-
ly. Essentially, the same relationships exist if
mean flows from January to June, inclusive,
are compared. This examination suggests that
reproductive success of woundfin (and round-
tail chub) in their only remaining habitat is
extremely poor when mean winter and spring
flows fall to about 100 cfs.
The drought of 1977, resulting in some of
the lowest flows on record in the Virgin Riv-
er, has permitted a significant insight into the
habitat requirements of the endangered na-
tive fishes of the river. It is apparent that
current utilization practices of the water re-
sources permit survival of the native fishes in
about 29 percent of their remaining potential
habitat. Intermittent flows coupled with
higher summer temperatures throughout the
remainder of the potential range (Schumann
1978, Lockhart 1979) make it unreliable as a
fish habitat. Within the remaining 29 percent
of the potential habitat, reproduction occurs
during years of normal flow, but is extremely
poor to absent during years of low flow. This
circumstance suggests that at present the
fishes are living in a habitat which has ex-
tremely little potential for further devel-
opment or alteration without adverse im-
pacts on the endangered species present.
Continued monitoring of reproductive suc-
cess and population structure under varying
conditions of stream flow will permit refine-
ment of flow requirements. It is apparent
that the roundtail chub is in an even more
precarious position than is the woundfin and
that both species require higher flows in
spring and winter than they do in summer.
Obviously, problems associated with the
80.
70
60
50
40
o upper river
D lower river
"T 1 1 1 1 • I
FALL SPRING FALL WINTER SPRING FALL WINTER
'73 '77 '77 '77 '78 '78 '78
Fig. 6. Comparison of mean size of woundfin in the upper and lower mainstream Virgin River 1973, 1977, 1978.
1979
The Endangered Species: A Symposium
49
effects of habitat modifications are complex,
often having been developing for more than
a century, and always difficult to quantify or
even specifically identify. The problems
identified and briefly examined here for the
Virgin River have numerous counterparts
throughout the West, as is obvious from the
fact that 97 percent of the western fishes list-
ed herein are on this list in part because of
the present or threatened destruction, modifi-
cation, or curtailment of their habitat or
range.
Disease and Parasitism
Wilson et al. (1966) and Seethaler (1978)
have suggested that parasitism may place sig-
nificant stress on western fishes being sub-
jected to other alterations in their environ-
ments. Examination of museum specimens of
Crenichthys boileiji collected since 1938, sup-
plemented by examination of both museum
specimens and individuals taken in the field
in 1965 and 1966, yields interesting insights
into responses to stress. Crenichthys baileyi
occurs in warm springs along the course of
the Pluvial White River of eastern Nevada.
During the early 1960s various exotic or non-
native species were established in some Cre-
nichthys habitats (Deacon et al. 1964, Hubbs
and Deacon 1964).
Figure 10 and Table 3 show the incidence
of parasitism by Lernea on Crenichthys bail-
eyi populations living in Crystal Spring and
in the warm headwaters springs of the
Moapa River from 1938 to 1966. All avail-
able data are presented in Table 3. Only data
resulting from an examination of 20 or more
individuals are plotted in Figures 10 and 11.
During this period no nonnative fish were es-
tablished in Crystal Spring. The poulation re-
mained abundant and virtually free of para-
sitism by Lernea.
In the headwaters of Moapa River, the
30
25
20
15
10
5
• — • 26 Nov. ■77(N=383)
O — o 28 Sept. '78(N=427)
20 30 40 50 60 70 80 90
Fig. 7. Length frequency of woundfin in the lower Virgin River, fall 1977 and fall 1978.
50
Great Basin Naturalist Memoirs
No. 3
23July
Lake Mead
Fig. 8. Dates of the first appearance of woundfin fry at various locations along the mainstream Virgin River dur-
ing summer 1977. Collections were made at all locations indicated at one- to two-week intervals until fry were
taken.
shortfin molly {Poecilia mexicana) was in-
troduced in the spring of 1963 (local testi-
mony). No mollies were taken in collections
made in March 1963, but they were present
in collections made on 12 October 1963
(Deacon et al. 1964). In addition, the mosqui-
tofish {Gambusia affinis) had been present in
the area since before 1938 (Miller and Alcorn
1946). While collections of C. baileyi made
during 1963 showed an increased incidence
of parasitism by Lernea, the fish population
remained abundant and the incidence of par-
asitism declined (Fig. 10). In addition, there
was a 5 percent incidence of parasitism in
1959 prior to introduction of mollies. Per-
haps the several spring sources in the head-
Table 3. Incidence of parasitism by Lernea on Crenichthys haileiji. N
one or more Learnea attached.
number examined. %
perc
ent with
Location
1938
N %
1940
N %
1941
N
1947
N
1948
N
1949
N
1950
N
Headwaters of
Moapa River
250
0
7
0
.\sh Spring
5
0
8
0
Crystal Spring
Hiko Spring
1576
0
6
0
Mormon Spring
Preston Town Spring
Preston Big Spring
0
1.58
0
11
0
11
0
106
0
64
0
20
0
64
0
83
0
54
0
52
0
58
17
0
0
1979
The Endangered Species: A Symposium
51
70.
•
• above narrows
0 below narrows
60.
50-
O
•
o
•
o
40
1
1 1 —
100 200 300 400 500 600 700 800 900 1000 1100
MEAN SPRING FLDW (CFS)
Fig. 9. Mean spring flows in Virgin River related to mean size of woundfin in the following fall. Data are from
1973, 1977, 1978. Mean spring flow is the average of the monthly means for April, May, and June.
waters were invaded by mollies at different
times; in any case, the Crenichthys popu-
lation did not appear to either sustain or re-
flect any permanent damage from parasitism.
Other populations for which historical data
are not so extensive but which, through 1966,
were not subjected to stress from nonnative
fishes occur at Mormon Spring and Preston
Big Spring (Table 3). In addition, while gup-
pies Poecilia reticulata have been in Preston
Town Spring since sometime before 1961
(Deacon et al. 1964), we have seen no in-
dication of parasitism by Lernea (Table 3). Of
course, the population was not examined im-
mediately after introduction of Poecilia.
Figure 11 illustrates changes in incidence
of parasitism by Lernea for populations
which became rare or extinct. In Ash Spring,
mosquitofish were not present in 1946 (Miller
and Alcorn 1946) but were present in 1959
(Miller and Hubbs 1960). In March 1963,
Poecilia was not present, but P. latipinna, P.
mexicana, and Cichlasoma nigrofasciatum
were present and breeding on 3 June 1964.
They have since remained abundant in Ash
Spring and its warm outflow stream. In-
cidence of parasitism by Lernea on C. baileyi
was significant for the first timme in 1964
and remained so in 1965. The C. baileyi pop-
ulation in this limnocrene declined in abun-
dance and remains extremely rare today. The
increase in parasitism closely followed in-
troduction of the exotics and was followed by
a dramatic decline in abundance of the na-
Table 3 continued.
1951 1954
N % N
1959
N
1960
N %
1961
N
1962
N
1963
N %
1964
N %
1965
N %
110
19
0
920
5.1
11 9
11
01
0
0
4
82
0
0
16 25
25 0
17 0
68
15
0 1259 0
253
0
90
10 224 9
5
0
2828 .04 356
0
69
0 313 20
27
59
159
0 1051 0
188
0
5
0 25 0
20
0 704 0
440
0
52
Great Basin Naturalist Memoirs
No. 3
tive fish population.
At Hiko Spring, no parasitism was evident
until 1965 (Fig. 11). Shortfin mollies (P. mexi-
cana), mosquitofish, and largemouth bass
were all absent from collections made at
Hiko Springs in June 1964. In January 1965 a
few mosquitofish were seen and one was col-
lected. In February 1965 both shortfin mol-
lies and largemouth bass were seen in the
limnocrene, and in March mollies began to
appear in the monthly collections. Both mol-
lies and mosquitofish increased in abundance
through 1965. Lernea first appeared on Cre-
nichthys in March 1965. Incidence of para-
sitism increased to February 1966, at which
time examination of the population was dis-
continued because numbers had declined too
low to permit continuation of the study. The
population was extinct before June 1967.
BioTic Interactions
Interactions of native western fishes with
introduced species have resulted in extensive
hybridization, especially in trout, plus vari-
ous kinds of competitive and predatory con-
sequences. One example which is especially
interesting, because it was replicated, oc-
curred in Manse Spring, Pahrump Valley,
Nye Co., Nevada. The endemic, and cur-
rently endangered, Pahrump killifish (Emp-
etrichthys latos latos) was restricted to the
single limnocrene which was approximately
triangular with maximum dimensions of
about 25 X 15 m. In November 1961 six
goldfish were introduced into the spring by
one of the farmhands. They had reproduced
by July 1962 and during that summer the
children on the farm removed most of the
30-,
25-
20.
E
o
o
CL
10.
■HaODD
•—^ headwaters of
Moapa River
Crystal Springs
Exotic Fish
Introduced
— I 1 1 1 1 1 1 —
1940 1945 1950 1955 I960 1965 1970
Fig. 10. Incidence of parasitism by Lernea on CrenichtJujs bailey i populations which remained abundant.
1979
The Endangered Species: A Symposium
53
submerged aquatic vegetation from the pond
to make a better swimming pool (Deacon et
al. 1964). The killifish population crashed
during the winter of 1962-63 to almost cer-
tainly fewer than 50 individuals (Fig. 12).
The population had recovered somewhat by
winter 1963 but appeared to be less abundant
through early 1965 than was the case prior to
introduction of goldfish.
In July 1967, Professors Carl L. Hubbs and
R. R. Miller and I, in cooperation with our
families and several students from UNLV and
ASU, attempted to remove all goldfish from
Manse Spring by trapping, seining, using
anesthetic, and, finally, dynamiting. All kill-
ifish captured were held in cages in a nearby
small spring and all goldfish were destroyed.
A total of 1239 killifish were captured and
returned. At least two adult goldfish eluded
us and spawned by the end of the summer.
The killifish population crashed as it had in
1963, reaching a low point of probably fewer
60
^
if)
25
20-
10-
•— • Ash Spring
Q-oHiko Spring
^ Exotic Fish Introduced
1940
1945
1950
1970
Fig. 11. Incidence of parasitism by Lernea on Crenichthys baileyi populations which became rare or extinct.
54
Great Basin Naturalist Memoirs
No. 3
than 50 individuals in July 1968. This low
population size persisted through January
1969 (Fig. 12), but by August 1971, when a
transplant was made into Corn Creek Spring,
the population had recovered significantly.
In August 1975, Manse Spring failed as a re-
sult of excessive pumping of groundwater in
the area (Soltz and Naiman 1978).
Prior to making the killifish transplant into
Corn Creek Spring the population of in-
troduced largemouth bass and mosquitofish
(Gambusia affinis) was removed. A few mos-
quitofish escaped the final poisoning efforts
in Corn Creek Spring, but by November
1973 the original stocking of 29 killifish had
built a population of about 1300. In addition,
mosquitofish had become extremely abun-
dant. By November 1974 approximately 250
killifish were estimated to occur in Corn
Creek Spring. The population had not in-
creased by July 1975. In April 1976, 165 kill-
ifish were removed from the spring and it
was poisoned in a second effort to remove ented by Soltz and Naiman (1978). Deacon
mosquitofish. The effort was successful and
killifish had built an estimated population of
2000 fish by November 1976 and 2500 by
October 1977.
These data show that on two occasions in
Manse Spring a population increase of gold-
fish was accompanied by a marked popu-
lation decline of Pahnimp killfish, and on one
occasion in Corn Creek Spring a population
increase of mosquitofish was accompanied by
a killifish population decline. A cause-effect
relationship is strongly suggested, perhaps re-
lating to competitive interactions of the
young or predation.
Restricted Range
While many western fishes have extremely
restricted ranges, none is so restricted or iso-
lated as the Devils Hole pupfish, Cyprinodon
diabolis. A discussion of the biology of this
species and description of its habitat are pres-
a TOTAL CAPTURE
0
1200 .
O MARK AND RECAPTURE A
6oldfls^ Reettoblilhod
1100.
• CATCH PER TRAP HOUR /
1000 .
/
900 .
/
800.
/
700.
/
/
600.
/
/
1
500 .
^^+-J^
\
1
400 .
Goldfith
\ /
/
300 .
INTRODUCTION
1/
% /
200 -
l\
Y
1^ /
100 .
[r
\ /
1961 1962 1963 1964 1965 1966 1967 1968
Fig. 12. Changes in population size of Pahnimp killifish 1961-1968.
1979
The Endangered Species: A Symposium
55
and Deacon (1979) provide a detailed de-
scription of fluctuations in population size
and probable causes for these fluctuations
through December 1976. Data on fluctua-
tions in population size presented here ex-
tend through December 1978 (Fig. 13, Table
4). Figure 13 illustrates the direct and
marked influence of relatively small changes
in water level in Devils Hole on minimum
population size of Cijprinodon diabolis. The
water levels indicated in Figure 13 refer to a
reference point established by USGS above
the maximum water level. Therefore, depth
of water in the habitat increases as the dis-
tance below the reference point (in feet) de-
creases. In addition, the water level shown is
actually the minimum level permitted by the
courts during the time indicated. The first
level indicated (3.9) represents the lowest wa-
ter level reached prior to intervention of the
courts. Water levels normally fluctuated
somewhat above the level indicated, but al-
most never below that level. Generally, wa-
ter levels were highest in winter and very
near the permissible minimum during the
summer irrigation season. This, of course, re-
flects the fact that the water level in Devils
Hole is directly and rapidly influenced by
pumping of groundwater nearby.
The somewhat erratic population fluctua-
tions in 1972 and 1973 reflect responses to
temporary management attempts as well as
to scouring floods v/hich occurred during this
period (Deacon and Deacon 1979). Once
550
500
450
400
350
300
250
200
150
100
1972 73 74 75 76 77 78
Fig. 13. Devils Hole pupfish population size compared to minimum water levels 1972-1978.
56
Great Basin Naturalist Memoirs
No. 3
some stability was achieved in water levels, it
became possible to attempt management of
water level to achieve a desired minimum an-
nual population size. The desired minimum
population size was established at 200 in an
effort to insure that the population would not
fall so low as to tend to accelerate toward ex-
tinction. The present court-mandated level of
2.7 appears to be just maintaining minimum
population size (Fig. 13, Table 4).
This example illustrates the direct and rap-
id impact on restricted native fishes which
can result from even modest developments
nearby. Often, as was true in this case, the
developer may be almost entirely unaware of
the consequences of his activities. For fishes
living in restricted environments, this lack of
awareness can mean extinction.
Discussion
It is apparent that the full variety of rea-
sons for becoming threatened are exemplified
among the endangered or threatened fishes of
the West. The legitimate question arising
from this and every consideration of endan-
gered species is "Why bother? What good
are they?" The answers to those questions, I
believe, must include at least two parts: (1)
because it is to our own self-interest to do so,
and (2) because our society's values, as ex-
pressed through federal law, require us to
"bother." The second answer has been and
will continue to be debated and perhaps
modified. The first is really the core of the
endangered species debate. The argument,
simplified, I believe, involves at least the fol-
lowing considerations. Because populations
are dependent upon and interact within eco-
systems, extinction is an indication of a signif-
icant change in the ecosystem— in general, a
reduced capability to support life or at least
to support diversity. The fact that an endan-
gered species is involved may, therefore, be
an indication that the long-term carrying ca-
pacity of an ecosystem may be exceeded (the
Table 4. Estimated population size of the Devils Hole pupfish [Cijprinodon diabolis) in Devils Hole, Nye County,
Nevada, 1972-1978. Estimates are the maximimi number of fish actually coimted visually during standardized at-
tempts at counting the entire population. Data prior to 4 June 1974 were taken by Dr. R. R. Miller and subsequently
by J. E. Deacon.
Population
Population
Population
Date
estimate
Date
estimate
Date
estimate
1972
1975
1977
6 April
127
22 Jan
208
20 Jan
324
2 June
248
20 Feb
159
24 Feb
276
28 July
286
18 Mar
148
24 Mar
198
27 Sept
191
10 Apr
158
5 May
210
14 Nov
199
19 May
201
6 June
221
16 June
262
27 June
359
1973
30 Jvily
278
11 July
330
10 Jan
252
20 Aug
294
15 Aug
553
20 Feb
191
30 Sept
260
26 Sept
490
28 Mar
208
21 Oct
279
28 Nov
381
12 June
184
25 Nov
261
.30 Aug
253
16 Dec
246
1978
6 Nov
244
16 Jan
296
1976
2 Mar
225
1974
17 Feb
228
16 Mar
219
5 Feb
163
3 Mar
180
24 Apr
223
29 Apr
143
30 Mar
181
19 May
242
5 June
2.39
27 Apr
195
19 June
274
11 Julv
250
18 May
203
20 Julv
326
22 Aug
286
22 June
.316
11 Aug
388
15 Sept
.302
2 Aug
.345
13 Sept
358
9 Oct
277
24 Sept
410
18 Oct
441
19 Nov
250
18 Oct
385
11 Dec
361
18 Dec
238
3 Dec
334
1979
The Endangered Species: A Symposium
57
argument of the canary in the coal mine).
Thus, it follows that if we are concerned
about the ability of our children to fimction
in the ecosystem in a manner at all com-
parable with our present functioning, it may
be important to maximize the survival of spe-
cies other than Homo sapiens who are also
dependent on that ecosystem.
Another major line of argument is the di-
versity-stability one (i.e., there appears to be
a tendency for more diverse ecosystems to be
more stable). Because more stable ecosystems
tend to permit coping with times of poor
productivity, it seems that enlightened self-
interest would dictate that we make efforts
to promote stability. Another cogent part of
this argument is the inverse relationship be-
tween diversity and energy flow (in molecu-
lar systems, ecological systems, and in organi-
zation of cities) described by Watt (1972,
1973). He pointed out that the principle ap-
pears to be true in societal organization to
the extent that in the U.S. we find fewer
book titles per capita than less industrialized
societies, as well as declining numbers of au-
tomobile and airplane manufacturers, in-
creasingly standardized foods in super-
markets and restaurants, symphony
orchestras almost restricting performances to
the work of eight men, difficulties with pub-
lishing innovative books or trying out in-
novative ideas, and declining numbers of spe-
cies (Watt 1972, 1973). In some ill-defined
way this general reduction in environmental
diversity seems to result in a search for re-
placement of the satisfaction or sensory stim-
ulation which it provided. Thus, we have sig-
nificant and expanding elements in our
society attempting to satisfy their senses
through membership in cults, sexual experi-
mentation, use of drugs and alcohol, etc.
Basically, it seems that as we manufacture a
more "efficient" society we increase its
energy flow while reducing its diversity. This
seems to result in a search for diversity by
the members of society. Perhaps the most
dramatic demonstration that environmental
stimulation derived from experiences with or
in nature is essential to modern man's feeling
of well-being comes from the successes real-
ized in the treatment of "hopeless" mental
cases (litis 1967). Dramatic improvements re-
sulted from taking these people on camping
trips. Many people obviously have expe-
rienced the tremendous release of tension
that can be felt when you "get away from it
all," or, to put it another way, when you
have an opportunity to become acquainted
with the diversity and sensory stimulation
available in nature. Finally, the availability
of genetic diversity in plants and animals as a
basis for producing new or better crops, med-
icines, and pharmaceuticals (Reisner 1978)
has been emphasized as one of the most com-
pelling arguments for saving species.
Thus, there are a number of biological rea-
sons to justify saving endangered species.
These usually have implications that extend
to other areas of human endeavor. If man's
uniqueness in fact is his knowledge of his
world, if Homo sapiens is the knowing one,
then each extinction diminishes man's capaci-
ty of know— and to that extent man's human-
ity. It seems to me that the Endangered Spe-
cies Act represents a society saying "This is
as far as we will go." The necessity of making
such a statement will always be questioned,
but it does represent an attempt at insuring
that our children on into many generations
will have available to them some of the hu-
manizing experiences that were available to
us.
Perhaps we have taken the position that
the extermination must stop because of our
general awareness that there is no other
choice. Human civilization has always had a
very nomadic character about it. The domi-
nant center of Western civilization has
shifted from the fertile crescent of Mesopo-
tamia to Egypt, Greece, Rome, Europe,
Great Britain, and the United States as envi-
ronmental overexploitation has forced (or
permitted) these nomadic wanderings. With
the entire planet occupied by civilized so-
cieties, there is no way to continue the wan-
derings of civilization. The last remnant of
the tendency appears to be exportation of the
environmental degradation required to sup-
port the kind of society we have created.
Thus, no longer does our civilization have its
primary impact confined to national bound-
aries. We find ourselves responsible for de-
struction of tropical rain forests, whales, pup-
fish, woundfin, and any number of other
worldwide resources, both renewable and
nonrenewable. A balance of payments deficit
58
Great Basin Naturalist Memoirs
No. 3
is clearly one serious and unacceptable con-
sequence, but it is completely overshadowed
by the rapid diminution of the world's ability
to support the biotic diversity so essential to
man's physical and mental well-being.
During this symposium Lovejoy (1979) has
provided a frightening description of the aw-
ful magnitude of the problem. Clements
(1979) has clearly shown that it is our own so-
ciety, not societies in the under-developed
countries of the tropics, that must be held
primarily responsible for such all-pervasive,
worldwide environmental degradation. Per-
haps an understanding of these important
facts will hasten the hard decisions which
must be made to apply the principles of the
Endangered Species Act on a worldwide
scale. Spencer (1979) provided extensive
documentation to show that the very difficult
and costly decisions essential to slowing the
rate of environmental degradation in the
United States are being made in some specif-
ic cases. His presentation is perhaps the most
encouraging evidence presented at the sym-
posium to indicate that there are forces at
work in our society which have a slim possi-
bility of forcing the significant shifts in so-
cietal values which Clements (1979) de-
scribed as essential if we are to prevent the
collapse of our system.
The answers to "Why save species?" are
many-faceted, almost always translate into
"Why save ecosystems?" and clearly demand
searching examination of human values. It
seems particularly powerful, therefore, to
find philosophers, theologians, and ecologists
converging on essentially the same answers to
these questions. Though ecologists tend to
understandably emphasize species and eco-
systems and theologians tend to emphasize
individuals and anthropocentricity, pretty
much the same conclusions emerge. The most
succinct and, to the Christian world, prob-
ably the most widely understandable con-
clusion we can arrive at was expressed by
Professor Hugh Nibley. In a 1978 essay exam-
ining man's relationship with his environ-
ment he said, "Man's dominion is a call to
service, not a license to exterminate."
Acknowledgments
Numerous people have assisted in the de-
velopment of data and ideas presented here-
in. To all I express sincere gratitude. James
D. Williams, Gail Kobetich, Thom Hardy,
and the American Fisheries Society Endan-
gered Species Committee were instrumental
in development of Tables 5 and 6. Jerry
Table 5. Described taxa of threatened freshwater fishes of western North America: 1979.
Present °
Historical
Common name
Scientific name
Status
threat
distribution
Trouts, family Salmonidae
Little Kern golden trout
Sahno aqttabonita whitei
Jordan
T
1,4
CA
Arizona trout
•Srt/mo apache Miller 1972
T
1,4
AZ
Lahontan cutthroat trout
Salmo clarki henshawi Gill
and Jordan
T
1,4
CA,NV,UT,WA
Colorado River cutthroat
Sulmo clarki pleuriticus
SC
1,4
CO,UT,WY
trout
Cope
Paiute cutthroat trout
Salmo clarki seleniris Snyder
T
1
CA
Greenback cutthroat
Sahno chirki stomias Cope
T
1,4
CO
trout
Utah cutthroat trout
Sahno clarki titali Suckley
T
1,4
UT,WY,NV
Rio Grande cutthroat
Salmo clarki virginalis
SC
1,4
CO,NM
trout
(Girard)
Gila trout
Salmo gilae Miller
T
1
NM,AZ
Sunapee trout
Salvelinus alpiniis atircolus
Bean
T
4
ME,NH,ID
Montana Arctic grayling
Thymalhis arcticus
T
1
MT
(stream form)
montanus (Pallas)
Mudminnows, family Umbridae
Olympic mudminnow Sovumhra hubhsi Schultz
SC
1979
The Endangered Species: A Symposium
59
Minnows, family Cyprinidae
Mexican stoneroller
Devils River minnow
Desert dace
Alvord chvib
Fish Creek Springs Tiii
chub
Independence Valley Tui
chub
Mohave Tui chub
Newark Valley Tui chub
Oregon Lakes Tui chub
Lahontan Tui chub
Owens Tui chub
Thicktail chub
Humpback chub
Bonytail
Gila chub
Chihuahua chub
Yaqui chub
Gila roundtail chub
Pahranagat roundtail
chub
Virgin River roundtail
chub
Oregon chub
Least chub
White River spinedace
Virgin spinedace
Big Spring spinedace
Little Colorado spinedace
Spikedace
Moapa dace
Yaqui Beautiful shiner
Rio Grande shiner
Proserpine shiner
Bluntnose shiner
Woundfin
Splittail
Colorado squawfish
Relict dace
Campostoma ornatum SC
Girard
Dionda diaboli Hubbs and T
Brown
Eremichthys dcros Hubbs T
and Miller
Gila ahordensis Hubbs and SC
Miller 1972
Gila bicolor euchila E
Hubbs and Miller 1972
Gila bicolor isolata Hubbs T
and Miller 1972
Gila bicolor mohavensis E
(Snyder)
Gila bicolor newarkensis SC
Hubbs and Miller 1972
Gila bicolor oregonensis SC
(Snyder)
Gila bicolor obesa (Girard) SC
Gila bicolor snyderi Miller E
1973
Gila crassicauda (Baird and E
Girard)
Gila cypha Miller E
Gila elegans Baird and E
Girard
Gila intermedia (Girard) SC
Gila nigrescens (Girard) E
Gila purpurea (Girard) E
Cw7fl robusta graluimi Baird T
and Girard
Gila robusta jordani Tanner E
Gila robusta semintida E
Cope
Hybopsis crameri Snyder SC
lotichthys phlegethontis T
(Cope)
Lcpidomeda albivallis T
Miller and Hubbs
Lepidomeda inollispinis T
inollispinis Miller and
Hubbs
Lepidomeda mollispinis E
pratensis Miller and Hubbs
Lepidomeda vittata Cope
Meda fulgida Girard
Moapa coriacea Hubbs and
Miller
Notropis formosus mearnsi
Snyder
Notropis jemezanus (Cope)
Notropis porserpinus
(Girard)
Notropis simiis (Cope)
Plagopteriis argentissim us
Cope
Pogonichtlnjs SC
macrolepidotus (Ayres)
Ptychocheilus luciiis Girard E
Rclictus solitarius Hubbs SC
and Miller 1972
1,3
1
L5
1
1,4,5
1,4,5
1,4
1,5
1
1
1,4,5
1,4,5
AZ,TX,(Mexico)
TX
NV
NV.OR
NV
NV
CA
NV
OR
NV
CA
CA
AZ,CO,UT,WY
AZ,CA,CO,NV,UT,WY
AZ,NM
NM,Mexico(Ch)
AZ,Mexico (So)
AZ,NM
NV
AZ,NV,UT
OR
UT
NV
Az,[/r
,vv
SC
T
E
1,4
1,3,4,5
AZ
AZ,NM
NV
SC
AZ (Mexico)
SC
T
L4
NM
TX
E
E
1,4
NM,TX
AZ,NV,UT
1,3,4
1
CA
AZ,CO,UT,CA,NM,NV,WY
NV
60
Great Basin Naturalist Memoirs
No. 3
Independence Valley
Rhinichthijs osculus
E
1,4,5
NV
speckled dace
lethoponis Hubbs and
Miller 1972
Ash Meadows speckled
Rhinichthijs oscidus
E
1,4
NV
dace
nevadensis Gilbert
Clover Valley speckled
Rhiniclitliijs oscidus
E
1,4,5
NV
dace
oligoponis Hubbs and Miller
1972
Kendall Warm Springs
Rhinichythijs oscidus
SC
5
WY
dace
thermalis Hubbs and
Kuehne
Moapa speckled dace
Rhinichthijs oscidus
moapae Williams 1978
T
1,3,4
NV
Loach minnow
Tiaroga cobitis Girard
SC
1,4
AZ,
Suckers, family Catostomidae
Yaqui sucker
White River desert
sucker
Webug sucker
Zuni bluehead sucker
Lost River sucker
Modoc sucker
Warner sucker
Shortnose sucker
Cui-ui
June sucker
Razorback sucker
Catostonius bernardini
Girard
Catostomus clarki
intermedins (Tanner)
Catostomus fecundus
Cope and Yarrow
Castostomus dicobohis
ijarrowi Cope
Catostomus luxatus
(Cope)
Castostomus microps
Rutter
Catostomus warneiensis
Snyder
Chasmistes brevirostris
Cope
Chasmistes ciijus Cope
Chasmistes liorus Jordan
Xyraiichen texanus
(Abbott)
Freshwater catfishes, family Ictaluridae
Yaqui catfish Ictahtrus pricei
(Rutter)
Widemouth blindcat Satan eurijstomus Hubbs
and Bailey
Toothless blindcat Trogloglanis pattersoni
Eigenmann
SC
T
SC
T
SC
E
E
T
E
SC
T
SC
T
T
Killifishes, family Cyprinodont
Railroad Valley
springfish
Leon Springs pupfish
Devils Hole pupfish
Comanche Springs
pupfish
Gila desert pupfish
Valley Amargosa pupfish
Ash Meadows Amargosa
pupfish
Warm Springs Amargosa
pupfish
Owens pupfish
White Sands pupfish
idae
Crenichthijs nevadae
Hubbs
Cyprinodon hovinus
Baird and Girard
Cyprinidon diabolis
Wales
Cyprinodon elegans
Baird and Girard
Cyprinodon macidarius
macularitis Baird and Girard
Cyprinodon nevadensis
amargosae Miller
Cyprinodon nevadensis
mioncclcs Miller
Cyprinodon nevadensis
pectorahs Miller
Cyprinodon radiosus Miller
Cyprinodon tularosa Miller
and Echelle 1975
1
1
1,4
1 c^jiu
1,4
1,4
1,4
1,4
1
1,4
1,4
1
1
1
SC
1
T
1,4,5
E
1,5
E
1
T
1,4
SC
1,4
T
1.4
E
1,4,5
E
1,4
SC
1,4,5
AZ (Mexico)
NV
UT
NM
CA,OR
CA
OR
CA,OR
NV
UT
AZ,CA,CO,NV,UT,WY
AZ (Mexico)
TX
TX
NV
TX
NV
TX
AZ, Mexico
CA
NV
NV
CA
NM
1979
The Endangered Species: A Symposium
61
Pahrunip killifish
Empetrichthijs latos lotos E
Miller
1,4,5
NV
Livebearers, faniilv Poecilidae
Amistad gambusia
Gambusia ajnistadensis
Peden 1973
E
1,5
TX
Big Bend gambusia
Gambusia gaigei Hubbs
E
1,5
TX
San Marcos gambusia
Gambusia georgei Hubbs
and Peden
E
1,4,5
TX
Clear Creek gambusia
Gambusia heterochir Hubbs
T
4
TX
Pecos gambusia
Gambusia nobilis (Baird
and Girard)
SC
1,4
TX,NM
Gila topminnow
Poeciliopsis occidentalis
(Baird and Girard)
E
1
AZ,NM
Sticklebacks, family Gasterosteidae
Unarmored threespine Gasterosteus aculeatus
stickleback williamsoni Girard
CA
Sunfishes, family Centrarchidae
Guadalupe bass
Micropterus treculi (Vaillant
and Bocourt)
Perches, family Percidae
SC
TX
Fountain darter
Etheostoma fonticola
(Jordan and Gilbert)
E
1,5
TX
Gobies, family Gobiidae
O'opu nakea
Awaous stamineus (Eydoux
and Souleyet)
SC
1,4
HI
Tidewater goby
Eucijclogobius newbernji
(Girard)
SC
1
CA
O'opu alamo'o
Lentipes concolor (Gill)
T
1
HI
O'opu nopili
Sicydium stimpsoni Gill
SC
1,4
HI
Sculpins, family Cottidae
Rougfi sculpin
Cottus asperrimus Butter
SC
1
CA
Utah Lake sculpin
Cottus echinatus Bailey and
Bond
E
1,4
UT
Shoshone sculpin
Cottus greenei (Gilbert and
Culver)
SC
I
ID
°1— Present or threatened destruction of habitat
2— Overutihzation
3— Disease
4— Hybridization, competition, exotic or translocated
5— Restricted natural range
Lockhart provided data on woundfin from
1973. Brian Wilson assisted greatly with de-
velopment of data on parasitism, which could
not have been accumulated without accessi-
bility to fish collections at the University of
Michigan Museum of Zoology (R. R. Miller),
the University of Nevada— Reno (Ira LaRi-
vers), and BYU (Dave White). Encour-
agement and approval of permits necessary
to the work has been provided by the Ne-
vada, Utah, and Arizona Game and Fish de-
partments. Stimulating discussions and
searching examination of values have been
provided by my family (Maxine, Dave, and
Jack and Cindy Williams) and by classes at
UNLV. Financial assistance for various as-
pects of work reported herein has been pro-
vided by the U.S. Fish and Wildlife Service,
National Park Service, National Science
Foundation, Sport Fishing Institute, U.S. Bu-
reau of Reclamation, University of Nevada-
Las Vegas, and the City of St. George, Utah,
through Vaughn Hansen Associates.
62
Great Basin Naturalist Memoirs
No. 3
Questions for Dr. Deacon
Once a species is on its way to recovery, how does
one determine what the population level or popu-
lation density would be for the species to be consid-
ered no longer in danger?
That is an extremely knotty problem. In the case of
the Devils Hole pupfish we were primarily con-
cerned with maintaining a large enough population
to prevent population instabilities that might tend
to accelerate the process of extinction. It is generally
understood that populations have a minimum size
below which they are unlikely to maintain viability.
Bob Miller at the University of Michigan did some
experimental rearing of other species of pupfish in
the 1940s and also performed a number of trans-
plants into springs devoid of fish. His work indicated
that experimental populations started with small
numbers of individuals tended to decline in abun-
dance after a few generations, sometimes to extinc-
tion. His numerous transplants of pupfish into other
natural waters were never successful if fewer than
200 individuals were transplanted, and in only two
instances were they successful when more than 200
individuals were transplanted. During the middle
1960s a graduate student of mine, Carol James (now
Ivy), did some work on the Devils Hole pupfish
which, in retrospect, indicated that its population
had probably never fallen below 200 individuals. Fi-
nally a transplant of 24 Devils Hole pupfish into an
artificial pond below Hoover Dam resulted in a pop-
ulation maximum of about 200 individuals, followed
by a decline to about 50 individuals. This pattern
suggested loss of viability may be occurring in the
transplanted population of Devils Hole pupfish. This
line of argimient was successful in establishing the
fact that it would be unacceptably dangerous to per-
mit the population of Devils Hole pupfish to fall be-
low 200 individuals. Once that point was established
it was not difficult to show, with four or five years of
monthly data on estimated population size, that a
water level of 2.7 was necessary to sustain a popu-
lation of no fewer than 200 individuals. These eco-
logical relationships are being reported in the sym-
posium volume on research in the national parks to
be published in 1979.
Q. At what level do you consider the population to not
be threatened?
Table 6. Undescribed taxa of threatened freshwater fishes of western North America: 1979.
Present °
Historical
Common name
Scientific name
Status
threat
distribution
Trouts, family Salmonidae
Alvord cutthroat trout
Salmo clarki ssp.
SC
1
OR
Humboldt cutthroat trout
Salmo clarki ssp.
SC
1
NV
Redband trout
Salmo sp.
SC
1,4
ca,or,id,nv
Minnows, family Cyprinidae
Catlow Tui chub
Gila bicolor ssp.
SC
1
OR
Sheldon Tui chub
Gila bicolor ssp.
SC
5
NV
Cowhead Lake Tui chub
Gila bicolor ssp.
SC
1
ca
Hutton Spring Tui chub
Gila bicolor ssp.
T
1
OR
Borax Lake chub
Gila sp.
T
1,5
OR
Foskett Spring speckled
Rhinichthys osctitus ssp.
T
1,5
OR
dace
Killifishes, family Cyprinodontidae
Preston White River Crenichthijs baileyi ssp.
springfish
Southern White River
springfish
Warm Springs White
River springfish
Devils River Conchos
pupfish
LeConte desert pupfish
Quitobaquito desert
pupfish
Crenichthijs baileyi ssp.
Crenichtliys baileyi ssp.
Cyprinodon eximius ssp.
Cyprinodon macidariiis ssp.
Cyprinodon macularius ssp.
T
4,5
NV
T
1,3,4
NV
SC
1,4,5
NV
T
1
TX
E
1,4
CA
SC
1,5
AZ
Sculpins, family Cottidae
Malheur mottled sculpin Cottus bairdi ssp.
SC
OR
• 1— Present or threatened destruction of habitat
2— Overutilization
3— Disease
4— Hybridization, competition, exotic or translocated
5— Restricted natural range
1979
The Endangered Species: A Symposium
63
A. I consider 200 piipfish to be one which puts the spe-
cies in approximately the position it was prior to the
appearance of man— not completely in that position,
but approximately. Now it's as threatened as it al-
ways was because of its restricted habitat, but it is
no more threatened because of man's activities.
Q. Would cleaning up the waters here in the West af-
fect the species population?
A. In those areas where pollution is a problem it cer-
tainly would. Almost anything that's proposed which
will modify habitats must be examined with respect
to the possibilities of adversely affecting species,
whether or not they are endangered. It doesn't nec-
essarily mean that, for instance, salinity control pro-
jects will affect the woundfin minnow. In fact, some
of my work has demonstrated that there is probably
a good opportunity to design salinity control projects
that will be unlikely to affect the mainstream fishes
of the Virgin River. That conclusion is expandable to
many other instances in the Southwest. The impor-
tant thing is to design projects that are compatible
with the habitat requirements of the species im-
pacted. In other words, cleaning up the waters of
the West could affect species in a number of ways,
both adversely and favorably.
Q. I'm not convinced that what you have said about the
proposal to not go ahead with the power plant in
Dixie is reasonable. The suggestion was that they di-
vert some of the water from the Virgin River into a
reservoir in Warner Valley and with that carry on
with their electrical work. Now, of course, it would
be a coal plant and this would be cooling water for
the hydro plant. What is the problem? How is it go-
ing to endanger that fish?
A. The question is how is the Warner Valley Project
likely to add to the threats to the woundfin minnow
and roundtail chub in the mainstream Virgin River.
Thanks very much for asking it, Vasco (Tanner). This
obviously is not a simple problem. The basic answer
I see is that the Warner Valley Project as projected
will alter the flows of the mainstream Virgin River.
The hydrologists point out that most of the water
will be taken during the winter and spring. Data I
presented here today indicate that during the low-
flow winter, spring, and summer of 1977, woundfin
and roimdtail chub reproduction was extremely low.
To the extent that the Warner Valley Project in-
creases the frequency with which flows similar to
1977 occur, that project will adversely impact the
endangered fishes living there. Essentially, the prob-
lem is that the data so far demonstrate that 1977,
which was a low-flow year, resulted in conditions in-
compatible with very much reproduction of those
two species. If you cut off that reproduction, you're
likely to cause an extinction. Certainly every time
you modify the flow regime of the Virgin River such
that the native fish populations living there miss a
year of reproduction, you're very demonstratively af-
fecting the capability of those species to maintain
themselves in the river. My conclusions here are
really based on the fact that we have demonstrated
very poor reproduction during a time which repre-
sents the kind of postproject flows we could expect.
Q. Of course I've seen that river fluctuate from great to
almost nothing, so naturally I don't see that there is
any justification for not going ahead with it. They're
going to get water from Warner Valley as well as
just divert a little from the Virgin River into the res-
ervoir.
A. The crux of the matter, I think, is what flows are
necessary to permit reproduction of the woundfin
minnow. The data I presented suggest that flows in
the neighborhood of 1(X) cubic feet per second are
necessary to permit reproduction of woundfin and
roundtail chub. In fact, there is some suggestion that
winter flows must be somewhat higher. If the Warn-
er Valley project doesn't reduce winter and spring
flows below about 110 cubic feet per second, then I
would say that there is likely to be no adverse im-
pact. On the other hand, if it does, and it was dem-
onstrated by the hydrological study that it would,
then it does represent an impact. I'm not saying you
shouldn't have the project. AH I am saying is, if you
reduce flows, you're going to impact the minnow
and the chub.
Q. Just a comment more than a question. I understand
the Warner Project during 1977 would not have
been allowed to divert because the water was so low
that project requirements would not have permited
diversion. The 1977 situation would not be repeated
unless there was another low-water year.
A. If that's the case, then I fail to see the basis for the
rather marked objections that have been raised to
the conclusions I have reached.
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fishes of Nevada, with a history of their in-
troduction. Trans. Am. Fish. Soc. 73:173-193.
Miller, R. R., and C. L. Hubbs. 1960. The spiny-rayed
cyprinid fishes (Plagopterini) of the Colorado
River system. Misc. Publ. Mus. Zool. Univ. Mich-
igan 115:1-39.
Minckley, W. L., and J. E. Deacon. 1968. South-
western fishes and the enigma of "endangered
species." Science. 159(3822):1424-1432.
NiBLEY, H. W. 1978. Nibley on the timely and the time-
less. Vol. 1 Religious Studies Monograph Series.
Religious Studies Center, Brigham Young Univ.,
Provo.
Peters, E. J. 1970. Changes with growth in selected
body proportions of the woundfin minnow (Pla-
gopterus argentissimiis) Cope: Cyprinidae. Un-
published thesis, Brigham Young Univ.
Reisner, M. 1978. What are species good for? Frontiers
42(4):24-27.
Schumann, P. B. 1978. Responses to temperature and
dissolved oxygen in the roundtail chub, Gila ro-
busta Baird and Girard. Unpublished thesis,
Univ. Nevada, Las Vegas.
Seethaler, K. 1978. Life history and ecology of the Col-
orado squawfish (Ptychocheilus luciiis) in the up-
per Colorado River Basin. Unpublished thesis,
Utah State Univ.
Smith, G. R. 1978. Biogeography of intermountain fish-
es. Great Basin Nat. Mem. 2:17-42.
SoLTZ, D. L., and R. J. Naiman. 1978. The natural his-
tory of native fishes in the Death Valley system.
Natural History Museum of Los Angeles County,
Science Series .30:1-76.
Spencer, D. A. 1979. The law and its economic impact.
Great Basin Nat. Mem. 3:25-34.
Trautman, M. B. 1957. The fishes of Ohio. Ohio State
Univ. Press, Columbus, Ohio.
Vaughn Hansen Assoc. 1977. Impact of the Warner
Valley water project on endangered fish of the
Virgin River. Vaughn Hansen Assoc, Salt Lake
City.
Watt, K. E. F. 1972. Man's efficient rush toward deadly
dullness. Nat. Hist. 81(2):74-82.
1973. A movable (disappearing) feast. Saturday
Review of the Sciences. l(l):56-57.
Williams, J. E. 1977. Adaptive responses of woundfin,
Plagopterus argentissimiis, and red shiner, Not-
ropis lutrensis, to a salt spring and their probable
effects on competition. Unpublished thesis, Univ.
Nevada, Las Vegas.
Wilson, B. L., J. E. Deacon, and W. G. Bradley. 1966.
Parasitism in the fishes of the Moapa River, Clark
County, Nevada. Trans. Calif.-Nev. Sec. Wildl.
Soc. pp. 12-23.
RARE AQUATIC INSECTS, OR HOW VALUABLE ARE BUGS?
Richard W. Bauinann'
Abstract.— Insects are an important element in the analysis of aquatic ecosystems, (1) because the limited dis-
persal abilities of many aquatic species means that they must make a living under existing conditions, and (2) be-
cause they are often sensitive to slight changes in water and stream quality, thus making excellent indicators of the
physical and chemical conditions in a system. Examples of rare, ecologically sensitive species are presented from the
Plecoptera, Ephemeroptera, and Trichoptera. Detailed studies of rare aquatic insect species should produce impor-
tant information on critical habitats that will be useful in the protection of endangered and threatened species in
other groups of animals and plants.
I use the term rare instead of endangered
or threatened, because no aquatic insects are
presently on the United States hst of endan-
gered fauna. The term is still relative,
though, because my experience indicates that
nearly every species can be plentiful if
sought at the right place at the right time.
Aquatic insects are useful to anyone study-
ing aquatic habitats because they usually
meet two criteria that are essential in assess-
ing aquatic systems. First, they often have
very limited dispersal abilities, which means
that they must stay and make a living under
existing conditions. Second, they are often
sensitive to slight changes in water quality, so
their presence or absence tells something
about the physical and chemical conditions in
the system.
Because the conditions present in a given
habitat determine which species can live
there, these organisms become living in-
dicators of water quality in aquatic ecosys-
tems. If these organisms are invertebrates
which exist at low levels in the food web, this
is an advantage. Invertebrates are easier to
study than are larger animals, because they
are more abundant and usually do not carry
the emotional stigma associated with large
vertebrates. They can also indicate adverse
habitat problems sooner, so that adjustments
can be made in water or stream quality be-
fore the top carnivores are severely affected.
During my studies of aquatic insects in
western North America, I have found that
the distribution patterns of certain species fit
nicely with a model of island biogeography.
Many stoneflies (Plecoptera) are mostly re-
stricted to pristine habitats characterized by
cold, clean continuously flowing streams at
high elevations or in special spring-fed habi-
tats. It thus follows that if the particular hab-
itat in which they occur is threatened, then
they almost automatically become rare and
may become extinct. Such species popu-
lations are often considered relicts of faunas
that were once more prevalent when more
ideal conditions occurred.
Studies on the stonefly genus Amphine-
mura (Baumann and Gaufin 1972, Bauman
1976) showed that it was a Palearctic genus
that had extended into the Neotropical
Realm and was still present in western North
America in limited relictual populations. Al-
though four species showed fairly wide distri-
bution patterns in the United States and Mex-
ico, three species were restricted to a single
mountain range. Amphinemura apache oc-
curred in the Chiricahua Mountains of Ari-
zona, A. reinerti, was limited to Sierra Potosi,
Mexico, and A. piiehhi was found in a moun-
tain drainage near Veracruz, Mexico.
These A7nphinemiira species are poor fliers
and almost need a water connection for dis-
persal. They live only in small streams that
flow all year around and are of high quality.
Their distribution patterns closely follow the
spruce-fir and high pine forests in the south-
western United States and Mexico. They are
'Monte L. Bean Life Science Museum and Department of Zoology, Brigham Young University, Prove, Utah 84602.
65
66
Great Basin Naturalist Memoirs
No. 3
thus good indicators of a special aquatic habi-
tat in western North America.
Another example of restricted distribution
in the stoneflies is Capnia lacustra Jewett,
which only occurs in Lake Tahoe. It carries
on its entire life cycle under water at depths
of 100-400 feet (Jewett 1963, 1965). Only
one other similar stonefly is known, and that
is the genus Baikaloperia (Zapekina-Dulkeit
and Zhiltzova 1973), in Lake Baikal, Siberia.
It is also wingless and possesses similar mor-
phological and ecological traits. It is not sur-
prising that these two deep, ancient lakes
contain similar rare species which evolved
under specialized conditions and will be lost
if their habitat is destroyed.
Much attention has been given to several
fish species that occur in the Colorado River
drainage such as the Humpback Chub, Ra-
zorback Sucker, and Colorado River Squaw-
fish. These fish developed through time in
another type of specialized habitat that ex-
cluded the salmonids and allowed other taxa
to radiate into the open niches. Invertebrate
species have also developed imder similar
conditions and several forms also occur in the
Colorado River drainage. Edmunds (1976)
lists several mayfly species which are rare
and restricted to the Colorado River drainage
and similar large warm rivers in western
North America.
The following are a few examples of rare
mayfly species and an indication of where
they occur in the United States: Analetris
eximia. Green River; Lachlonia saskotchewa-
nensis. Green, Colorado, and White Rivers;
Anepeorus rusticus. Green River; Home-
oneuria sp. Escalante and Colorado Rivers.
These invertebrate species provide additional
evidence that our large, warm, western rivers
contain animal species that have adapted to
special conditions critical for their survival.
Caddisflies or the Trichoptera are inter-
esting insects that occur in a wide variety of
aquatic habitats. Most are good fliers and dis-
tribute freely, but many species are restricted
to a certain habitat because of the larval re-
quirements.
Wiggins (1977) published an outstanding
work that makes it possible for any trained
biologist to classify caddisfly larvae to genus.
Thus it provides another tool for evaluating
habitats using aquatic insects. A few inter-
esting examples and their special habitat re-
quirements are: Goriella baiimanni, organic
ooze in spring seeps; Psychronia costalis,
small meadow streams above 8,000 feet; Des-
mona bethida, small spring streams.
The number of rare aquatic insects is quite
large because of the number of different
aquatic habitats available and the ability of
insects to fit into relatively small niches with-
in those habitats. This is actually a positive
value, however, because it allows the re-
searcher to more closely understand the eco-
system since it can be divided into smaller
parts.
Two final examples of rare insects that
have very specific habitat requirements are
the met-winged midges and the water penny
beetles.
Net-winged midges are flies which have
become adapted to living in torrenticolus
habitats. The larvae are greatly modified into
chitonlike organisms that attach themselves
to the substrate by sucking discs. They live
only in clean, cold, well-aerated waters
which have a stable, smooth-grained sub-
strate. Thus they can be excellent indicators
of these habitat conditions that occur at falls
and quick-flowing mountain torrents. Hogue
(1973) lists several Blephariceridae species
that are presently known only from a single
locality or mountain range. This is not simply
an artifact of incomplete collecting, but a re-
sult of poor adult dispersal ability plus the
very specialized habitat requirements of the
larvae noted earlier.
Water penny beetles have an adult stage
tliat looks like a terrestrial beetle but a larval
stage that is highly modifed for life on the
bottom of streams. The larva is greatly flat-
tened so that the head and appendages are
completely hidden under the thoracic and
abdominal sclerites. The single eastern spe-
cies Psephenus herricki is rather widely dis-
tributed, but the five known western species
have very restricted distributions (Brown and
Murvosh 1974). Two species, P. montanus
(White Mountains, Arizona) and P. arizo-
nensis (Chiricahua Mountains, Arizona) have
very limited areas of occurrence. An inter-
esting note is that this type of limited distri-
bution pattern is also exhibited by several
species in the Plecoptera, Trichoptera, and
Ephemeroptera.
1979
The Endangered Species: A Symposium
67
Many more examples of "rare" aquatic in-
sects could be given which probably fit into
the endangered or threatened categories as
presently understood. They are exciting to
me for pure scientific studies of zoogeo-
graphy and phylogeny. However, I feel that
the real value is not simply to say "I found
another rare creature," but instead to make
us more sensitive about the critical habitat
conditions which produced these rare species.
Insects tend to be more abundant and are
tlius easier to study without affecting popu-
lation dynamics. They are also usually more
economical and easier to sample because
tliey are less mobile and can be effectively
studied by fewer people with less sophis-
ticated equipment.
On the other hand, politicians and business
people may question the value of an insect.
Who cares about bugs? How much is a bug
really worth?
This problem can be illustrated by an in-
cident with which I was involved while at
the Smithsonian Institution. Soon after my
arrival in Washington, D.C., I was asked to
look through the aquatic insects for which I
was responsible as curator and add^any spe-
cies to a list of organisms that could be con-
sidered both rare and restricted to the Chesa-
peake Bay area. In Ross and Ricker (1971) I
foimd a stonefly species, AUocapnio zekia,
that was known only from the Zekiah
Swamp, La Plata, Charles County, Maryland.
I added it to the list and forgot about it.
About three years later, I received a tele-
phone call from a man who wanted to know
all about the "Zekiah Stonefly." At first I did
not know what he was talking about, but
when he mentioned the Chesapeake Bay spe-
cies list, I remembered. I did some quick re-
search and indicated that the species was
based on a single, male holotype and might
possibly be a synonym of a widespread east-
ern species. He nearly exploded when I re-
ported this to him, because, he said, that "Ze-
kiah Stonefly" was holding up the
construction of a water plant in a nearby
community and was costing a lot of people a
lot of time and money.
In summary, it is important to understand
our special environmental problems here in
North America. If this can be better facil-
itated by using aquatic insects, then we
should place renewed emphasis on studies of
them. We must, however, be aware of the
fact that people in general do not imderstand
the scientific value of insects and might react
poorly to "bugs" being used to justify the
preservation and conservation of special eco-
systems. However, scientific investigations of
high quality must utilize all possible avenues
of investigation if problems are to be solved
with a minimum expenditure of time, effort,
and resources.
It is also important that we do not attempt
to overstate the value of "rare" species as
habitat indicators. Some states, for example,
have placed entire orders such as the Plecop-
tera (stoneflies) and Ephemeroptera (may-
flies) on lists or proclamations and have di-
luted their value. When this is done it
becomes difficult to study these organisms,
because of the problem involved in obtaining
permits and permission to collect specimens.
Collecting alone will probably never se-
riously harm an aquatic insect population, as
has occurred with many butterfly species, but
habitat manipulation will.
Literature Cited
Baumann, R. W. 1976. Amphinemuia reinerti, a new
stonefly from northern Mexico (Plecoptera:
Neniouridae). Southwest. Natur. 20:517-521.
Baumann, R. W., and a. R. Gaufin. 1972. The Am-
phinemiira ventista complex of western North
America (Plecoptera: Neniouridae). Natur. Hist.
Mus. Los Angeles Co. Contr. Sci. 226:1-16.
Brown, H. P., and C. M. Murvosh. 1974. A revision of
the genus Psephenus (Water-Penny Beetles) of
the United States and Canada (Coleoptera, Dryo-
poidea, Psephenidae). Trans. Amer. Entomol.
Soc. 100: 289-340.
Edmunds, G. F., Jr., S. L. Jensen, and L. Berner. 1976.
The mayflies of North and Central America.
Univ. Minnesota Press, Minneapolis.
HoGUE, C. L. 197.3. The net-winged midges or Blepha-
riceridae of California. Bull. Calif. Insect Sur. 15:
1-83.
Jewett, S. G., Jr. 1963. A stonefly aquatic in the adult
stage. Science 139: 484-485.
1965. Four new stoneflies from California and
Oregon. Pan-Pacific Entomol. 41: 5-9.
Ross, H. H., and W. E. Ricker. 1971. The clas.sification,
evolution, and dispersal of the winter stonefly
genus A//oca^ni«. Illinois Biol. Monog. 45: 1-166.
Wi(;r,iNS, G. B. 1977. Larvae of the North American
caddisfly genera (Trichoptera). Univ. Toronto
Press, Toronto, 4 dp.
Zapekina-Dulkeit, J. I., and L. A. Zhiltzova. 1973. A
new genus of stoneflies (Plecoptera) from Lake
Baikal. Entomol. Obozv. 52: 340-345.
ENDANGERED AND THREATENED PLANTS OF UTAH: A CASE STUDY
Stanley L. Welsh'
,\bstract.— Endangered and threatened plants of Utah are evaluated as to their distribution in phytogeographic
subdivisions, substrates, plant communities, elevations, and geological strata. The phytogeographic subunits were
partitioned and comparisons made of distribution as outlined for the parameters cited above. A predictive model is
suggested based on the nonrandom distribution of endemic plant species.
The Endangered Species Act of 1973, Pub-
lic Law 93-205 (as ammended 1978), was an
outgrowth of decades of concern regarding
the future of that portion of our heritage of
hving things, which, by the nature of their
distributional patterns, could most easily be
eradicated as man pressed to exploit the re-
sources of the earth, both finite and renew-
able. The act dictated an orderly process for
development of lists of endangered and
tlireatened species, defined terminology, and
provided for development of criteria for de-
termining candidate species.
Plants are the mantle of the land, nour-
ishers of life's feast, holders of soil, suppliers
of construction materials, of medicines, and
of other substances too numerable to men-
tion. They provide the basis of all life on
earth, save some few living things which are
capable of chemosynthetic utilization of
energy. This fact and the list of materials that
flows from plants need not be mentioned.
Yet, the spread of mankind over the face of
the earth, his development of agriculture,
and, more especially perhaps, his devel-
opment and spread of an industrial society
with its great demands on space and mate-
rials has resulted in a direct competition for
the space that was, or is, occupied by the in-
digenous flora of the earth.
The clearing of agricultural land for plan-
ting of crop plants, as selected from that in-
digenous genetic stock available as portions
of the total flora, was possibly the beginning
of the role of mankind as a major agent for
reduction of plant species. Even those from
which the crop plants were developed were
not spared from destruction or modification.
Agriculture is, nevertheless, a more ef-
ficient means for the production of biological
materials that can be consumed by man and
by his livestock than from the previously em-
ployed methods of gathering and hunting.
Industrialization merely speeded the pro-
cess by which agricultural lands could be
cleared of native plants and those lands then
maintained in single crop cultures. With in-
dustrialization came the explosion of de-
mands for resources of many kinds: ferrous
and nonferrous metals, chemical compounds
of all kinds, sand and gravel, coal and oil,
uranium, and other naturally occurring mate-
rials.
The mantle of the land gave way as each
new source was discovered. Roads were cut
through the vegetation. Quarries, open pit
mines, portals, corridors, industrial plant
sites, pipelines, villages, towns, cities, gar-
bage dumps, litter, and other features of civ-
ilization were placed atop the shrinking veg-
etation.
Into the vast array of plant species
marched also an infinitesimally small cadre of
'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Prove, Utah 84602.
70
Great Basin Naturalist Memoirs
No. 3
persons determined to know about the plants
themselves— to name them, to describe them,
to plot where they grew, and to recognize
that there is an intrinsic value in each plant
species, no matter how insignificant it might
be considered. Botanists they were called,
whether by training or by inclination they ar-
rived at a point where plants become their
pursuit. At first, all botanists were tax-
onomists. Later, not even all taxonomists
were taxonomists.
Late in the human story the taxonomists
began to catalogue the vegetation of the
earth. Systematic surveys of vegetation and
collections of plant species began in earnest
only in the eighteenth century, in North
America not until the nineteenth century,
and in Utah the main thrust did not come un-
til the twentieth century.
By the beginning of the third decade of the
present century, the common plant species
and their general areas of growth were well
known. The work of the various government
surveys and of pioneer botanists had pene-
trated even to some of the most remote re-
gions of western North America. Discovered
were some of the most rare of species, but
others remained undiscovered.
Cognizant of the increasing demands of a
growing population and an expanding civ-
ilization, botanists, always too few for the
task, were hard pressed to survey all of the
remote regions in a systematic manner. Col-
lections were taken in a haphazard way. A
trip to the hot desert in springtime, another
to the cool mountains in midsummer, and by
autumn the enthusiasm for collecting was cut
short, too often by the need for gainful em-
ployment—because botanists could seldom be
gainfully employed as botanists.
As the search areas narrowed, and as col-
lections were taken in a more systematic
manner, the number of known narrowly re-
stricted plants increased proportionally. A
still finer search may yet yield many addi-
tional narrowly adapted endemics. They are
plants of all elevational ranges, but they are
most common in highly specialized habitats,
those which are likely to be occupied by oth-
er narrowly restricted plants also.
Often the species belong to difficult or to
purportedly difficult taxonomic groups, such
as Astragalus, Eriogonum, Erigeron, and oth-
ers. Few people have taken the time to un-
derstand these complex assemblages, or to
even collect and attempt to identify them.
Fortunately, monographers have examined
many of the problem genera and have clari-
fied the nature of taxonomic limits, often on
the basis of very limited materials.
Passages of the Endangered Species Act
found botanists in most regions of the United
States ill prepared to provide definitive infor-
mation regarding candidate plant taxa, which
had been included in the act mainly as an af-
terthought. Despite the lack of specific infor-
mation, the act called for the secretary of the
Smithsonian Institution in Washington, D.C.,
to report to Congress within one year on all
of the "species of plants which are now or
may become endangered or threatened" in
the United States (Section 12, Public Law 23-
205). In December 1974, the secretary of the
Smithsonian Institution, S. Dillon Ripley,
submitted a "Report on Endangered and
Threatened Plant Species of the United
States" to Congress.
That report formed the basis of the 1 July
1975 Federal Register (Vol. 40, No. 124:
27824-27924), which contained a review of
the endangered and threatened plant species.
The number of species assigned to those cate-
gories for the twelve western states (exclusive
of Hawaii) is presented in Table 1. That pre-
Table 1. Number of species reviewed as endangered
and threatened in 1975 and proposed as endangered in
1976, in twelve western states.
Date
1975
1976
Status
Status
STATE
E
T
E
Alaska
9
21
6
Arizona
65
106
66
California
236
412
286
Colorado
23
17
32
Idaho
21
41
21
Montana
2
8
3
Nevada
43
84
50
New Mexico
15
26
20
Oregon
43
135
51
Utah
56
101
66
Washington
16
72
18
Wyoming
3
18
8
Total
582
Grant total =
1623
1041
627
1979
The Endangered Species: A Symposium
71
liminary list of 1975 was based on the best in-
formation available to scientists at the Smith-
sonian Institution working in collaboration
with those from the Department of the Inte-
rior. The lists were reviewed by selected spe-
cialists and botanists at a workshop held at
the Smithsonian in September 1974.
That the 1975 lists were preliminary is to
be found in the differences in numbers of en-
dangered species published in the Federal
Register (Vol. 41, No. 117: 24524-24572)
published on 16 June 1976 (Table. 1). Even in
such states as California, with its formidable
niunber of qualified professional taxonomists
and amateurs, the number of endangered
plant candidates increased significantly be-
tween 1975 and 1976. No such comparable
list is available for candidate threatened
plants, but some of the increase in endan-
gered species is represented in change of stat-
us from threatened to endangered (Kartesz
and Kartesz 1977).
Impetus for acquisition of knowledge of
rare plant species was generated by the lists
of 1975 and 1976, and by the policy of active
search for information required by govern-
mental agencies, which was built into the act.
Funds were forthcoming from various federal
agencies to make determinations of range,
habitat, condition, impacts, and potential im-
pacts, and for other information on the candi-
date species. Rule making was entered into
by the U.S. Fish and Wildlife Service, De-
partment of the Interior, and, at present,
some 20 species of plants have been deter-
mined as endangered or threatened. Two of
these. Astragalus perianus Barneby and Pha-
celia argillacea Atwood, are from Utah (see
Federal Registers Vol. 40, No. 81:
17910-17916, and Vols. 43, No. 189:
44810-44812, respectively). The former is
listed as threatened, and the latter is listed as
endangered.
Impacts of the act have been widespread.
It has been subjected to political and emo-
tional, as well as to scientific, evaluations.
The act has been modified to some extent as
a result, but those evaluations are not the
basis of this contribution. Rather, I intend to
pursue the developmental basis of informa-
tion dealing with endangered and threatened
species, and to outline one basis of the nature
of those critical plants.
Biology of Endangered and Threatened
Species
The biology of endangered and threatened
species in Utah is, with few exceptions, the
biology of narrowly restricted endemics.
Therein lies the basis for disparity between
lists and category representations. The
amount and quality of botanical knowledge
of common species is seldom sufficient to al-
low more than generalizations; that for rare
species is likely to be lacking altogether. The
task of surveying vast areas for narrowly re-
stricting plants is a huge one, carried out in
the past largely by individuals with much de-
votion and little financing.
Too, the fact that a plant is an endemic
and is rare has often been considered as evi-
dence of endangerment. Lists are replete
with such examples, but studies have in-
dicated that rare plants might not be endan-
gered or threatened, and that plants thought
to be rare were in fact relatively common
and widely distributed. For a plant to be a
candidate for inclusion on final lists of endan-
gered and threatened plant species, it must
have endangerment, both quality and quan-
tity, clearly demonstrated.
Contemporary studies are under way to
aid the Department of the Interior with deci-
sions necessary for final rule making. Studies
of distribution, population numbers, degrees
of endangerment, and many other facets are
being undertaken, which will lead to devel-
opment of information summaries of all spe-
cies which have been reviewed, proposed, or
recommended.
Much information has already been
gleaned from the specimens extant in her-
baria. For the purpose of this paper, endan-
gered and threatened plant species from Utah
will be used to illustrate the contemporary
knowledge of status of those species, and to
provide the model for a case study of the na-
ture of those species.
A list of endemic and rare plants of Utah
was prepared by Welsh, Atwood, and Reveal
(1975). In that publication, some 382 vascular
plant taxa were considered, with 66 regarded
as endangered, 198 as threatened, 7 extinct,
and 20 extirpated. Only 225 species were
considered to be endemic to Utah. The num-
bers are not comparable to those published in
72
Great Basin Naturalist Memoirs
No. 3
the Federal Registers due to consideration of
species with broad distribution, a portion of
which includes Utah, within the threatened
and endangered categories. In later com-
putations the number of endemic species is
cited as 239 (Table 2). Welsh (1978) pub-
lished a reevaluation of the endangered and
threatened plants of Utah, in which some 53
species are regarded as endangered and 99 as
threatened. Numbers in this latter pub-
lication are not comparable to those of the
Federal Register lists due to deletions and ad-
ditions.
That the biology of endangered and threat-
ened species is that of restricted local endem-
ics is found in the nonrandom distribution of
those species. Utah can be divided into elev-
en phytogeographic subunits, each topo-
graphically, geologically, and phytologically
different (Table 2). The numbers of endan-
gered and threatened plants is approximately
proportional to the number of endemic spe-
cies in each phytogeographic subunit. En-
demics constitute 27 percent of the total for
the Navajo Basin; endangered and threatened
plant species of that basin make up 28 per-
cent of the total for the state. Proportions are
similar for Plateau, Tavaputs, Uinta Basin,
and all Qher phytogeographical regions. Ap-
proximately 64 percent of all endemic spe-
cies in these areas are considered as endan-
gered or threatened. It is axiomatic that
endemics should constitute the endangered
and threatened candidates when the small
areas occupied by them are considered.
Endangered and threatened species of the
Navajo Basin and Plateau subunits constitute
half of the total number for Utah. Other im-
portant regions include the Uinta Basin (13
percent), Great Basin (13 percent), and Mo-
have (14 percent). The remaining areas in-
clude only 11 percent of the species on can-
didate lists in total.
In general, endangered and threatened
plant taxa in Utah occupy harsh substrates
which are perceived by man as barren or
nearly barren of vegetation. Hence, these
critical species tend to occur in areas where
there is little competition. Survival of the
species depends on maintenance of the habi-
tat in a condition wherein other species do
not become competitive. Protection, as here-
in conceived, involves guarantees against
man-caused destruction of habitat. Natural
changes should not be treated as endan-
germent.
Phytogeographic Subunits and
Endangered and Threatened Species
The distribution of critical species does not
appear to be at random on the substrates
available, and those substrates which support
these species are not occupied uniformly.
Rather, specific portions of apparently sim-
ilar substrates are occupied, while others are
not. Clays and other fine-textured coUuvial or
aeolian materials and limestones are the most
commonly occupied substrates (Table 3). To-
gether, they form the substrates of 81 percent
Table 2. Comparison of endemic, endangered, and threatened plant taxa by phytogeographic subdivision within
Utah.
Phytogeographic unit
Endemi
ics
Endangered
Number Percent N
Threatened Total I
umber Percent Number
• &t
Number P«
jrcent
Percent
7
3
1
1
1
1
65
27
17
32
25
25
42
28
48
20
11
21
22
22
33
22
3
1
2
4
2
1
25
10
8
15
12
12
20
13
4
0
4
4
4
3
3
1
2
2
2
1
23
10
1
2
6
6
7
5
35
15
7
13
13
13
20
13
26
11
'
13
14
14
21
14
Colorado canyons
Navajo Basin
Plateau
Tavaputs
Uinta Basin
Uinta Mts.
Wyoming
Wasatch Mts.
Great Basin
Mohave
Pine Valley Mts.
239
53
100
99
1979
The Endangered Species: A Symposium
73
Table 3. Endangered and threatened plant species arranged by substrate within Utah.
Endangered Threatened
Total
Substrate Number Percent Number Percent
Number Percent
Clay, silt, mud
Sand
Gravel
Igneous gravel
Limestone
Talus
Loam-humus
Water
Unknown
L5
2
9
8
101
100
of the taxa on critical lists. Water-washed al-
luvial deposits tend to support a greater
plant cover than do the in situ substrates or
those deposited by wind. Hence, probably
because of the greater amount of cover and
competition, the vast areas covered by water-
borne alluvial deposits mainly lack critical
species. The exceptions include some gravel
deposits, especially those derived from ig-
neous extrusive or intrusive stocks.
Critical species are present in several of
the major vegetative types within the state.
As with other criteria, the taxa are not ran-
domly distributed in all of the plant commu-
nities. Pinyon-juniper, desert shrub, warm
desert shrub, and salt desert shrub vegetative
types bear 65 percent of all endangered and
threatened candidates (Table 4). These are
communities which lie within a low precipi-
tation regime, wherein edaphic features are
not insulated from plants by well-developed
soil horizons or by organic matter within the
soils. Edaphic features are the main con-
trolling factors within low-elevation plant
communities.
Even where plants of a critical nature are
present within a community which tends to
occupy most of the surface area, and where
soils are well developed, thus preventing di-
rect edaphic control, the endemic species are
found mainly in clearings, along bluff mar-
gins, on ridge tops, and on other poorly vege-
tated micro-habitats.
Adiabatic and lapse rate differentials are
reflected in elevational differences. High ele-
vation areas are cooler and receive propor-
Table 4. Endangered and threatened plant species arranged by plant community (the plant communities in ap-
proximate order by elevation).
Community
Alpine
Spruce-fir
Aspen
Mountain brush
Ponderosa pine
Pinyon-juniper
Sagebrush
Desert shrub
Warm desert shrub
Salt desert shnib
Hanging garden
Aquatic
Other
Unknown
Endar
igered
Percent
Threatened
Total
Number
Number
Percent
Number
Percent
5
5
5
3
3
6
14
14
17
11
2
4
1
1
3
2
1
2
1
1
2
1
2
4
7
7
9
6
17
.32
27
27
44
29
2
4
3
3
5
3
12
23
4
4
16
11
5
9
9
9
14
9
4
8
22
22
26
17
1
1
1
1
1
1
1
1
1
1
1
1
4
8
4
4
8
5
53
101
99
99
152
100
74
Great Basin Naturalist Memoirs
No. 3
tionally greater amounts of precipitation, re-
sulting in production of mesophytic plant
communities in those sites. Plant species of a
critical nature are mainly xerophytes, regard-
less of the community type within which
they occur. The large portion of species,
some 60 percent of those designated as en-
dangered or threatened, exist below the 6000
foot (1930 m) contour (Table 5). Possibly the
reason for the great number of species at the
lower elevations is due to the proportionally
greater number of sites in arid lands which
are open to colonization.
Chemical and water relations of substrates
are closely allied to geological strata. Eda-
phic control by geological formations is
greatest in areas where the strata are ex-
posed. Layers of alluvium, which represent
mixtures of materials from different sources,
tend to insulate vegetation which grows on
that alluvium from the chemical and water
relations peculiarities of the individual stra-
tum per se. Soil development reinforces sepa-
ration of parent materials from plants.
Hence, geological control of vegetative cover
is greatest at lower elevations, where strata
of many kinds are exposed over vast reaches.
Soils as such are poorly developed or nonex-
istant due to low rainfall and the corollary
lack of leaching of soluble salts.
There are regions at moderate to high ele-
vations where edaphic factors of geological
strata are controlling due to peculiarities of
topography and geomorphology. Cliff faces
and breaks at the margins of plateaus and
ridge crests are examples of such places. In
others, substrates which are very acidic or
basic, as in some igneous or limestone strata,
tend not to be insulated due to lack of
growth of dense vegetation. Plant species of a
critical nature occur on a series of geological
strata ranging in age from Quaternary to Pre-
cambrian (Table 6).
There does not appear to be any particular
stratum which bears a disproportionately
large number of endangered or threatened
species. The largest number is found on Qua-
ternary alluvia, mainly on dunes or stabilized
dune sand and on residual accumulations on
the formations from which they were pro-
duced. Even this small number represents
only 17 percent of the included species.
Dunes are open habitats. They are mesophyt-
ic sites in otherwise arid lands. They repre-
sent an anomaly wherein competition is low,
but where water is relatively abundant and
available.
If mudstone, siltstone, and shale strata are
considered collectively, some 37 percent of
the species reside on them. Limestone or oth-
er highly calciferous formations, such as
Flagstaff, Wasatch, and the Carboniferous
strata, provide substrates for 17 percent of
the total plant species. Sandstone and con-
glomeritic formations account for only 10
percent of the taxa.
Partitioning of the phytogeographic sub-
divisions demonstrates differences and sim-
ilarities in areas of distribution, and in the
control of that distribution. Disparity in geo-
logical strata is obvious from one subunit to
the next, and potential substrates differ be-
cause of the different kinds of strata avail-
able. The Paleozoic strata of the Great Basin
and of the Wasatch Mountains present an en-
tirely different array than do the Uinta
Mountains, Uinta Basin, Navajo Basin, and
Mohave subunits. Plant communities reflect
those substrate differences, often in subtle
ways. Additionally, the phytogeographic sub-
imits are topographic features whose defini-
tions are tied to elevation.
Despite the problems associated with com-
parison, and the obvious differences— which
should not require discussion— an analysis of
the various phytogeographic subunits will be
Table 5. Endangered and threatened plant species arranged by elevation stratification.
Elevation
<6000 feet (1830 m)
6000-9000 feet (1830-2745 m)
>9000 feet (2745 m)
Unknown
Endan
gered
Percent
Threa
tened
Total
Number
Number
Percent
Number
Percent
34
64
58
58
92
61
13
24
30
30
43
28
3
6
10
10
13
9
3
6
1
1
4
3
53
100
152
100
1979 The Endangered Species: A Symposium 75
instructive in attempts at management of limestone. The other phytogeographic sub-
lands in the respective areas as regards en- vmits bear so few species as to not demon-
dangered and threatened species. The total strate trends.
numbers of species in a given subunit might When plant communities are compared for
be indicative of trends (Tables 7, 8, 9, and each of the phytogeographic subunits, it is
10). clear that pinyon-juniper and the various
Summaries of species number and percent- kinds of desert shrub communities support
ages for substrates in each of the phytogra- most of the endangered and threatened plant
phic subunits demonstrates similarities be- species in the Navajo Basin, Uinta Basin, and
tween the Navajo Basin, Uinta Basin, and Mohave subunits in Utah (Table 8). Spruce-
Mohave subunits (Table 7). In each of these, fir, ponderosa pine, and pinyon-juniper com-
clay, mud, silt, and sand constitute the sub- munities are the sites of occurrence of some
strates of more than 85 percent of all critical 71 percent of the critical species in the
plant species. Plateau subunit differs in bear- Plateau subunit. Alpine and spruce-fir are the
ing more than 50 percent of the included main communities of those species in the
species on limestone, and with igneous Uinta and Wasatch mountains,
gravels being second with 18 percent. Pat- The Navajo, Uinta, Great Basin, and the
terns in the Great Basin are obscure, with no Mohave subunits bear 80 to 100 percent of
single substrate supporting more than 25 per- the species below 6000 feet in elevation. In
cent of the species. Six of the seven species Plateau, Tavaputs, Uinta Mountains, and
from the Wasatch Mountains are known from Wasatch Mountains all species are above the
Table 6. Geological strata as substrates of endangered and threatened Utah plant species (Note: species were
assigned to only one stratum, the major one, even if they occurred on more than one. Strata without numbers of
species, indicated by a dash, are known to support critical species; those not marked are not known to support
them.)
Strata
Quaternary
Flagstaff
Green River
Bald Knoll
Wasatch
Duchesne River
Tertiary Igneous
Kaiparowits
VVahweap
Straight Cliffs
Mancos Shale
Tropic Shale
Mowry
Arapien
Cedar Mt.
Morrison
Entrada
Carmel
Navajo
Wingate
Chinle
Moenkopi
Cutler
Cedar Mesa
Paradox
Carboniferous
Precambrian
Unknown
Threatened
Endangered
Number Percent
Total
Number
Percent
Number
Percent
18
18
8
1.5
26
17
2
4
2
1
.3
3
10
19
13
8
9
9
4
8
13
8
5
5
5
3
9
9
1
2
10
7
2
2
2
1
1
1
1
1
5
5
4
8
9
6
1
1
1
1
2
4
3
1
2
1
4
1
4
1
4
1
3
1
—
—
1
1
1
2
2
1
2
3
2
1
1
3
6
4
3
5
5
3
6
8
5
1
1
1
1
7
1
7
1
5
1
9
2
12
2
8
1
1
1
1
2
2
1
12
12
1
2
13
8
.3
3
3
2
7
7
4
8
12
8
99
99
53
99
152
99
76
Great Basin Naturalist Memoirs
No. 3
Table 7. Substrates of endangered and threatened plant species by phytogeographic subdivision in Utah.
Substrate
Colorado
Navajo
Uinta
Canyons
Basin
Ni
Plateau
Basin
Number Percent
Number Pe
rcent
umber
Percent
Number Percent
23
55
2
6
16 80
1 100
16
38
4
12
3 15
1 5
11
2
6
17
18
52
1
2
1
2
2
1
1
4
3
3
Clay, silt, mud
Sand
Gravel
Igneous gravel
Limestone
Talus
Loam-humus
Water
Unknown
TOTAL
100
2b
6000-foot contour (Table 9). Part of the ex-
planation for this correlation is based on the
definition of the subunits. The basins are
mainly below 6000 feet in elevation, and the
mountains are mainly above that figure.
Similarities of geological formations in
chemical and physical structure seem to be
more important than the geological strata by
themselves. Cutler, Moenkopi, Chinle, Car-
mel, phases of the Entrada, Morrison, Ara-
pien. Tropic Shale, Mancos Shale, and Du-
chesne River formations tend to resemble
each other texturally, and in having high
amounts of soluble salts. Each of these sup-
port one or more of the endangered or
threatened species, some of which might be
expected on others of those formations also.
Indeed, some do occur on more than one for-
mation, even though Table 10 is presented
with only the major formation that serves as
substrate represented. Differences and sim-
ilarities between the subunits of the state are
obvious. Geological strata in subunits of the
Colorado drainage system tend to be most
similar, but even in those there is a tendency
for plants to react differentially, despite the
similarities of stratigraphy.
Predictive Capability
Because of the nonrandom distribution of
narrowly restricted species in Utah, it is pos-
sible to prepare a model with predictive ca-
pability which will aid in the search for these
critical plants. The model, a sample of which
is presented in Table 11, is based on deduc-
Table 8. Plant communities of endangered and threatened plant species by phytogeographic subdivision in Utah.
Plant Comm.
Alpine
Spruce-fir
.\spen
Mountain Brush
Ponderosa pine
Pinyon-juniper
Sagebrush
Desert shrub
Warm desert shnib
Salt desert shnib
Hanging garden
Aquatic
Other
Unknown
Colorado
Canyons
Navajo
Basin
Plateau
Tavaputs
Number Percent Number Percent Number Percent Number Percent
12 2 6
9 27
1 3
100
99
100
1979
The Endangered Species: A Symposium
77
Table 7 continued.
Uinta
Wasatch
Great
Mts.
Wyoming
Mts
Basin
Moh
lave
Tavaputs
Number Percent
Number Percent
Number '.
Percent
Number
Percent
Number
Percent
Number Percent
5
25
9
43
2 100
2 100
4
2
1
20
10
5
9
43
6
86
5
25
2
10
1 25
3 75
1
5
1
14
2
10
1
5
4 100
2 100
7
100
20
100
21
101
2 100
tions derived from the nature of the distribu-
tion of those species evakiated above. The
reasoning behind the model is based on the
unequal occurrence of the species with re-
gard to several parameters. The probabilitv
of occurrence is determined by use of a nu-
merically weighted system in which the pa-
rameters are given a value of zero, one, or
two as indicated by the known presence of
the species on specific portions of the state.
For example, most of the species of restricted
plants occur on the finely textured soils, the
next highest proportion on dunes, in situ
sand, and limestone, and the lowest on soils
consisting of gravel, talus, loam, and humus.
Hence, these substrate types are rated as two,
one, and zero, respectively.
The example outlined in Table 11 is sug-
gested for the state, but more finely parti-
tioned models are suggested for each of the
phytogeographic subunits. It would not be
logical to apply a high numerical weighting
to elevations below 6000 feet for areas within
the Plateau phytogeographic subunit where
practically all the known critically restricted
plants are above that elevation.
Despite the usefulness of the suggested
model, and its modifications, it is suggested
that the model should be used as a planning
tool only. There is no substitute for on-
ground inspection and the collection of the
general flora to provide information on ac-
tual presence of plant species. Wherever pos-
sible such on-site investigations should pro-
vide herbarium materials for deposit in
herbaria, taken in such manner as not to con-
Table 8 continued.
Uinta
Basin
N:
Uir
Ml
ita
:s.
Wyoming
Wasatch
Mts.
Great
Basin
Mohave
Number
Pe
ircent
umber
Percent
Number Percent
Number Percent
Ni
jmber
Percent
Number
Percent
3
1
60
20
1
5
14
71
8
40
1
20
1 50
1
14
1
3
5
15
1
7
5
.33
9
3
45
15
1 50
10
4
2
50
20
10
2
9
2
10
43
10
20
100
7
100
2 100
7
99
20
100
21
101
78 Great Basin Naturalist Memoirs No. 3
Table 9. Elevation of endangered and threatened plant species by phytogeographic subdivision in Utah.
Colorado
Canyons
Navajo
Basin
Plateau
Tavaputs
Elevation
Number Percent Number Percent Number Percent Number Percent
< 6000 ft.
6000-9000 ft
> 9000 ft.
Unknown
100
100
33
100
100
stitute a threat in and of itself. This will guar-
antee that information gained in field surveys
will not be lost in the files of agencies and in-
dustries attempting to work on the lands of
the state.
Perspective on the
Endangered Species Act
Value judgements as to the role of plants of
limited distribution have not stopped, slowed
down, or even modified the course of human
expansion through all of history until now.
Tlie present society has asked whether plant
species should be eradicated as a part of the
common good of our civilization. Value is a
time-oriented function; that considered as
valueless today might be judged as very valu-
able in the future. Numerous examples of
minerals are known which support this obser-
vation. Plants have been surveyed many
Table 10. Geologic strata serving as substrates of threatened plant species by phytogeographic subdivision in
Strata
Quaternary
Flagstaff
Green River
Bald Knoll
Duchesne R.
Wasatch
Tertiary
Kaiparowits
Wahweap
Straight Cliffs
Mancos
Tropic
Dakota
Mowry
Arapien
Cedar Mt.
Morrison
Entrada
Carmel
Navajo
Wingate
Chinle
Moenkopi
Cutler
Cedar Mesa
Paradox
Carboniferous
Precambrian
Unknown
Colorado
Canyons
Navajo
Basin
Plateau
Tavaputs
Number Percent
Number Percent
Number
Percent
Number Percent
4
10
3
2
9
6
2 100
2
5
14
7
2
42
21
6
1
8
3
1
2
20
67
2
1 100
1
1
3
3
4
3
2
2
2
2
7
7
10
7
5
5
1
2
3
6
3
7
2
6
1 100
41
98
a3
99
2 100
1979
The Endangered Species: A Symposium
79
Table 9 continued.
Uinta
Basin
Uinta
Mts.
Wyoming
Number Percent
Wasatch
Mts.
Great
Basin
Mohave
Number Percent
Number Percent
Number '.
Percent
Number
Percent
Ni
umber Percent
20 100
20 100
2
3
5
40
60
100
1
1
2
50
50
100
5
2
7
71
29
100
17
1
2
20
85
5
10
100
19 90
2 10
21 100
times for sources of biologically active mate-
rials, e.g., alkaloids, vitamins, hormones, won-
der drugs, and cortical steroids. Now they are
again being surveyed for antitumor agents.
Common plants are being surveyed first, pri-
marily because of their ready availability.
The rare plants will be reviewed as material
becomes available. It would be a tragic irony
if the best anticarcinogenic agent should be
discovered in the leaves of a herbarium speci-
men of a species which had just become ex-
tinct.
Extinction occurs as a function of natural
forces, or as a function of man-caused factors.
The former is selective in reducing popu-
lations of living things. The latter is non-
selective.
The reasons for extinction of narrowly re-
stricted plants on the same outcrop might in-
volve loss of a pollinator for one species, in-
Table 10 continued.
Utah (Note: Only the major formation is indicated where plants occupy more than one.)
Uinta
Basin
Uinta
Mts.
Wyoming
Wasatch
Mts.
Great
Basin
Mohave
Number Percent Number Percent Number Percent Number Percent Number Percent Number Percent
50
35
20
20 100
20
100
100
100
25
15
100
5
101
80
Great Basin Naturalist Memoirs
No. 3
festation by insects or disease for another. A
construction project might cause wholesale
extirpation by removal of the entire commu-
nity. The rate of man-caused extinction far
exceeds the natural rate. Thus, extinction
caused by man is not a part of the natural
scheme.
The Endangered Species Act of 1973 made
it possible for future generations to be in-
volved in the value-oriented decisions. The
act provides an advocate for generations yet
unborn.
Genetic pathways are, despite all of the
possibilities, essentially one-way streets. The
route by which a species is formed is as im-
portant as the end result. The reconstitution
of the pathway requires the same criteria as
were present in the past, a functional impos-
sibility to recreate. Thus, the loss of any spe-
cies terminates a line which cannot be re-
formed. And, once gone, the question of
value to mankind is deprived of practical sig-
nificance.
The reason most of the proposed endan-
gered and threatened plants are considered
thusly is because the known populations are
small and exist in very limited areas. Average
distributional densities of one endangered
species to each two or three thousand square
kilometers, and of threatened species to val-
ues of roughly half that figure, give an ap-
proximation of their true paucity. Further,
only a very small part of the total land sur-
face is involved.
Distribution of rare species is not equal, as
has been discussed above. Certain areas ap-
pear to lack them altogether, while other
areas support concentrations of several spe-
cies. Unless a specific mineral to be exploited
is located within one outcrop which supports
one or more species, or unless the area to be
occupied by a particular development is
large, there is no reason why modern expan-
sion should impress any of the currently
known endangered or threatened species.
Even in these two exceptional instances there
is no real reason to displace indigenous en-
dangered and threatened species; the best site
for industrial development is not always the
only good alternative.
Thus, if developers, and if the govern-
mental agencies which control development
on federal lands, follow the requirements as
set forth in the act, there is little question
that many, if not all, of the plant species
which are ultimately determined as endan-
gered or threatened can persist in perpetuity.
The question of value of these plants is not
an issue; the areas occupied by these plants
can be avoided.
Literature Cited
Kartesz, J. T., AND R. Kartesz. 1977. The biota of
North America. Part 1. Vascular plants. Volume
I— Rare plants. Bonac, Pittsburg, Pa. 361 pp.
Welsh, S. L. 1978. Endangered and threatened plants of
Utah; A reevaluation. Great Basin Nat. 38: 1-18.
1978. Status reports of endangered and threat-
ened plants of Utah. U.S. Fish and Wildlife Ser-
vice (unpubl. ms.).
Welsh, S. L., N. D. Atwood, and J. L. Reveal. 1975.
Endangered, threatened, extinct, and rare or re-
stricted Utah vascular plants. Great Basin Nat.
35: 327-376.
Welsh, S. L., and K. Thorne. 1979. Identification man-
ual of endangered and threatened plants of Utah.
U.S. Fish and Wildlife Service publication. 399
pp.
Table 11. Outline of a predictive model for establishing priority areas for study of endangered and threatened
plants of Utah.
Numerical
weighting
Substrate
Community
Elevation
Geology
Phyto Subunit
0
Gravel, talus,
loam, humus
Other
9000
Other
Colorado
Canyons,
Wyoming,
Pine Valley
1
Dunes, in situ
sand, limestone
Spruce-fir
Ponderosa
6000-90(X)
Sandstone,
in situ sand
and limestone
Wasatch Mts.,
Uinta Mts.,
Tavaputs PI.,
Plateau
2
Clay, silt, mud
Pj-Des
Sh variations
6000
Shale mud
and siltstone
Navajo, Uinta,
Mohave, Great
Basin
MANAGEMENT PROGRAMS FOR PLANTS ON FEDERAL LANDS
Diiaiie Atvvood'
.\bstr\ct.— The plant phase of the Endangered Species Program is discussed from the point of view of a profes-
sional botanist in government service. Some of the new amendments are also discussed from a botanical standpoint.
Federal agency programs and policies in the western United States are briefly reviewed. The strength of the Endan-
gered Species Program is dependent upon input from qualified professional biologists in and out of government ser-
vice. Some of the problems encountered in the program are outlined.
The comments I would like to make today
are based on my experience with government
agencies over the past several years. I do not
speak as a representative of any government
agency, although I have had experience with
the Bureau of Land Management (BLM),
Fish and Wildlife Service (FWS), and the
Forest Service (FS). First of all, let me point
out that professional people who work for
government agencies have a very frustrating
and difficult task. They want to get on with
the job that should and could be done, but
cannot because of regulations, policies, and
conflicts with the management and planning
staff. There is a communication gap between
professionals and managers and planners that
needs to be bridged somehow. Until recently,
some of these agencies were strictly manage-
ment and planning oriented. Passage of new
federal laws and regulations, such as the En-
dangered Species Act (ESA), created a need
for these agencies to hire professionals with
specialized training. It must be recognized
that managers and planners have a difficult
job making the proper decisions for the best
uses of our natural resources and still be in
time with the multiple use concept. Our job
as professionals is to supply managers and
planners with sufficient data on any given
problem or project, as it relates to our area of
responsibility and expertise, so they can eval-
uate the pros and cons and in turn make the
proper decisions. The active support of the
Endangered Species Program (ESP) varies
from agency to agency and from state to
state within a given agency. For example.
California has an excellent and effective
Threatened and Endangered (T/E) plant pro-
gram at both the state and federal level. Both
state and federal agencies there have active,
qualified botanists. Additional professionals
outside of government have also taken an ac-
tive interest in the ESP.
In discussing various topics with the par-
ticipants of this symposium, I was impressed
with the need to clarify the responsibilities of
the different agencies that participate in the
ESP. As most of you know, the Fish and
Wildlife Service has taken the lead in this
program for terrestrial species and has the re-
sponsibility for developing and implementing
regulations to guide other federal agencies
and the states in meeting the purpose and in-
tent of the ESP. To accomplish this task they
have published guidelines to implement the
Convention on International Trade for En-
dangered Species of Fauna and Flora, prohi-
bitions on certain uses of endangered and
threatened plants, criteria for determining
critical habitat, and the Inner Agency Coop-
erative Section 7 Regulations. In addition,
the Fish and Wildlife Service has the respon-
sibility for the consultation process, as re-
quired by the Section 7 Regulations, and the
listing and delisting processes. To most of us
here the listing process is the activity the
Fish and Wildlife Service should be moving
forward with most rapidly. However, they
have a disproportionate share of the work
load and budgeting restrictions have been
placed on them. Other major responsibilities
of the Fish and Wildlife Service include law
'Uinta National Forest, Forest Service, U.S.D.A., 88 West 100 North, Pn
81
82
Great Basin Naturalist Memoirs
No. 3
enforcement, land acquisition, cooperative
agreements with states, and development of
recovery plans and/or teams. The new
amendments to the ESA require some
changes in the program. One of the new
amendments now allows for the acquisition
of land for plants. Prior to these new amend-
ments. Section 5 of the act, regarding land
acquisition, was only for wildlife species or
plants officially listed and concluded in ap-
pendices to the convention. This new amend-
ment is a breakthrough for plants. As I un-
derstand it, the Forest Service, as well as the
Department of the Interior, can now acquire
land for plants. Formerly, the Department of
the Interior was the only federal department
that could acquire land. Additionally, there is
the new requirement for development of re-
covery plans for all officially listed endan-
gered and threatened species. In Utah we
have two plant species officially listed that
are either on or adjacent to Forest Service
Lands. We will be developing additional
background data for use in these two recov-
ery plans. I have two slides on them. The first
is of Astragalus perianus, which is endemic
to two locations in the central part of Utah at
high elevations. The species was originally
collected in 1905 by some of our early botan-
ists, but was not rediscovered until 1976. The
other species is Phacelia argiUacea, which is
endemic to the Green River Shale formation
along the railroad right-of-way in Spanish
Fork Canyon. This is the only existing popu-
lation that we know of, and only nine indi-
vidual plants exist, based on counts made in
1978. In view of the restricted nature of this
species, the Fish and Wildlife Service will
place this one high on their priority list for
development of a recovery plan.
The various phases of the program that the
Fish and Wildlife Service are trying to devel-
op and implement directly affect the activi-
ties of other federal agencies, particularly
land-managing agencies such as the BLM,
Forest Service, and National Park Service. As
most of you know, the Forest Service and
BLM are trying to develop active programs.
The National Park Service apparently takes
the position that threatened and endangered
species in the parks are already protected
and that they don't really need to do any-
thing. However, as Stan Welsh pointed out.
the influx of people into these areas does
have a detrimental effect on many T/E spe-
cies that exist there. Some of the other
agencies who have no lands to manage but
have an impact on endangered and threat-
ened species are the Bureau of Reclamation,
the Soil Conservation Service, and the Navy,
Army, and Air Force. For example, projects
with which the Bureau of Reclamation is in-
volved will destroy habitat. There is, there-
fore, a direct conflict with the purpose and
intent of the ESA when endangered or
threatened species are impacted by those
projects. Some of those agencies are making
no effort to determine the impact their proj-
ects have on these species. We as profes-
sionals, I feel, have the responsibility to be-
come aware of their projects and to help
provide these agencies with data and exper-
tise. The trend among federal agencies is to
solicit information and public opinion on var-
ious projects. How many of you are respond-
ing?
To comply with the objectives and policies
of the Endangered Species Program, the
BLM, Forest Service, and Fish and Wildlife
Service have developed the following policy
to insure protection for T/E species prior to
official listing and protection under the En-
dangered Species Act. These agencies are
considering all species that are likely to be-
come endangered or threatened as though
they are already officially listed to insure
their actions do not jeopardize the existence
of these species or modify their critical habi-
tats. The degree of implementation varies
within each agency from state to state and
even within a given state. The strength of the
program at these levels is dependent upon
the professionals available to insure program
development. There are very few plant tax-
onomists in government to help guide the
program. Therefore, the scientific commu-
nity must become more involved if we are to
achieve a realistic program. The benefits of
such a policy are fourfold: (1) protection of
sensitive species prior to listing, which can
and will meet the purpose and intent of the
1973 ESA, thereby (2) preventing the need
for official listing of many T/E Species, (3)
resulting in fewer legal restrictions and more
management options for agencies, and (4)
creating more benefits to the species and
1979
The Endangered Species: A Symposium
83
project development. A major concern of
federal agencies is to meet the requirements
of Section 7 of the ESA, which reads
The Secretary shall review all programs administered
by him and utilize such programs in hirtherance of the
purpose of this act. All other federal departments and
agencies shall in consultation with and with the assist-
ance of the Secretary utilize their authorities in furthe-
rance of the purposes of this act by carrying out pro-
grams for the conservation of endangered species and
threatened species listed pursuant to Section 4 of this
act and by taking such action as necessary to insure that
actions authorized, fimded, or carried out by them do
not jeopardize the continued existance of such endan-
gered and threatened species or result in the destruction
or modification of habitat of species which is deter-
mined by the Secretary after a consultation with the af-
fected states to be critical.
However, the overriding concern is to
meet the purpose and policy of the ESA,
"... to provide a means whereby ecosystems
upon which endangered species and threat-
ened species depend may be conserved, to
provide a program for the conservation of
such endangered species and threatened spe-
cies . . . [and the] . . . policy of Congress that
all Federal departments and agencies shall
seek to conserve endangered species and
threatened species and shall utilize their au-
thorities in furtherance of the purposes of
this Act."
It is my interpretation that the intent in
the purpose and policy of the act is to con-
serve and protect species likely to be endan-
gered or threatened with extinction in the
foreseeable future. John Spinks has indicated
that the Fish and Wildlife Service will only
be able to list 20 to 30 species of plants in
1979. This is less than 1 percent of the 1785
proposed species. The Forest Service and Bu-
reau of Land Management policy, if it is en-
forced, will provide the necessary protection
for species which would otherwise become
extinct due to the slow listing process. Fur-
thermore, such a policy will minimize the
need for official listing under the ESA.
Two other major problems in the plant
program come to mind: (1) a lack of data on
candidate and proposed species, and (2) in-
adequate lists of threatened and endangered
plants. The latter is a result of insufficient
data. Therefore, we must emphasize the need
for additional inventories to determine the
range of these species, their habitats, infor-
mation on population biology, threats to
their survival, and management problems.
Presently a lack of funds is the biggest ob-
stacle in developing an efficient data base.
Some contracts have been let, and the cur-
rent trend is to acquire these data through
new contracts. Once we determine the loca-
tions of the T/E plants, we have to go back
to these specific locations and obtain suf-
ficient data for use in management programs.
My assigned topic today was on management
programs for plants on federal lands. The fact
is federal agencies have formulated few or no
management programs for most plants be-
cause we are in the inventory stage at the
present time. We do have sufficient data on
some species to make recommendations on
listing or delisting from candidate and pro-
posed lists and establish monitoring studies
for others. The purpose of these monitoring
studies is to acquire additional data on the
status of the populations, their trends, condi-
tion of the habitat, and the biological needs
of the species to develop realistic manage-
ment programs for their protection and re-
covery, if possible.
California has an active program that
places them well ahead of other states. Most
of the other western states are developing
programs. Much of this effort is from the pro-
fessional and private sectors and the rest
from federal and state agencies. The state
government, in most states, has shown the
least interest and, in general, leans more to
development. Four federal agencies will issue
contracts for plant inventories in Utah this
year. It is hoped these studies will be con-
ducted by qualified professionals. In addition,
we have established coordinating committees
for state, federal, professional, and amateur
botanists in most of the western states to
avoid duplication of effort and coordinate
program activities. Botanists in Utah have
now established a Utah Native Plant Society.
One function of the society will be to help
implement a T/E plant program for the
state. It is hoped our program will be as suc-
cessful as that in California. We solicit your
membership if you have an interest in the na-
tive flora of Utah.
Most federal agencies do not employ plant
taxonomists. Fortunately, they do have some
biologists with sufficient interest and back-
ground to help develop a plant program. The
84
Great Basin Naturalist Memoirs
No. 3
Forest Service will hold training sessions for
existing range and wildlife staff to familiarize
them with T/E species in their areas of re-
sponsibility. As a zone botanist, I am respon-
sible for the Forest Service T/E plant pro-
gram in Utah and Nevada. Within this area
there will be from two to three hundred proj-
ects requiring site inspections for T/E plants.
With the current level of funding and avail-
able staff we can expect to look at only 10
percent of these projects until more funding
and personnel are available. We have, there-
fore, prioritized the species and areas to be
worked on. The initial effort is on critically
endangered species. The following slides il-
lustrate some of these. The first is Phacelia
argillacea, which, as I mentioned earlier, is
officially listed. Next is Townsendia oprica. It
is known only from two populations and, as
Stan Welsh indicated, one population had
been destroyed by a gypsum operation. Only
one population remains. Aictomecon humilis
is restricted to the Moencopi formation in
Washington County, Utah. It is more com-
mon, but the impacts to the area are so se-
vere that immediate listing is necessary to in-
sure protection.
Government-funded inventory contracts
have resulted in range extensions for many of
the proposed species, as well as the discovery
of new species. Psoralea pariensis is a species
described in 1975. Just a few years ago. Pri-
mula specuicola was known only from a few
collections along the Colorado River drain-
age system. Recent studies, as a result of the
ESA, have provided many new locations and,
even though from 30 to 40 percent of the
habitat has been inundateed by Lake Powell,
official listing is not necessary.
In the West, much of the land is adminis-
tered by federal agencies. Table 1 illustrates
the number of acres under Forest Service,
Bureau of Land Management, National Park
Service, and Fish and Wildlife control. Prob-
ably 5 percent or less of all these acres will
constitute critical habitat requiring pro-
tection for T/E plant species. However, until
our inventories are complete, we will not
know where that 5 percent of the
632,992,185 acres is. Again we must use a
priority system for inventories, based on the
minimal data available. To show another
relationship, I have outlined the number of
candidate, possibly extinct, proposed and of-
ficially listed species by state. Currently
there are 15 plant species officially listed.
More than half of them occur in California.
The new amendments now include plants
in Section 6 of the act under Cooperative
Agreements. Table 2 outlines the status of co-
operative agreements with states prior to the
new amendments. Even though plants were
not included in Section 6 of the act, original-
ly four states submitted proposals to the FWS
requesting funds for plants. Naturally, none
have qualified. However, Utah submitted
Table 1. Number of acres and T/E plants in the western United States.
(1975 FR)
(1976 FR)
Number of
(1975 FR)
Number of
BLM
USES
NPS
FWS
Candidate
Possibly
Proposed
Officially listed plants'
admin.
admin.
admin.
admin.
threatened
extinct
endangered
State
acres
acres
acres
acres
species
species
species
Threatened Endangered
Alaska
272,673,528
20,622,014
7,306,037
22,236,273
21
6
Arizona
12,596,043
11,219,839
1,629,943
877,200
106
5
65
California
15,577,909
20,327,515
4,258,2123
68,944
415
28
282
13
Colorado
8,354,671
13,773,966
535,050
51,947
18
3
32
Idaho
11,985,079
20,342,387
86,425
40,944
41
21
Montana
8,141,498
16,767,962
1,159,505
539,340
8
1
3
Nevada
48,373,664
5,112,7.55
262,321
2,202,045
85
6
48
New Mexico
12,956,665
9,106,299
241,621
316,183
26
3
19
Oregon
15,739,792
15,486,367
160,881
466,011
135
2
51
Utah
22,641,037
7,990,271
888,936
97,944
102
7
65
1 1
Washington
306,692
9,069,287
1,801,428
128,466
72
2
19
Wyoming
17,536,891
8,679,047
2,310,653
20,640,923
44,787
7,070,084
18
1,047
3
60
8
619
Totals
446,883,469
158,397,709
1 14
•22 species of the T/E plants have been officially listed.
1979
The Endangered Species: A Symposium
85
their proposal in June 1977 for both plants
and animals and is close to qualifying. This is
based on my conversation with the Washing-
ton office of the FWS. Some states have heri-
tage programs, and research natural area
councils that have been extremely helpful in
developing plant programs for the respective
states.
Your attendance at this symposium is evi-
dence of the interest shared by many Ameri-
cans in preserving our unique flora and
fauna. We can have the necessary devel-
opment to sustain us and still preserve these
valuable resources by having an open mind
to the problems at hand. Let's help close the
communication gap between scientists, envi-
ronmentalists, and politicians.
Questions for Dr. Atwood
Q. Is the listing of these taxa being coordinated because
there are so few that are going to be hsted? There
are strategies whereby protecting one species in a
very interesting habitat would preserve maybe four
or five others in the same area.
A. It's my understanding that the Fish and Wildlife
Service in-house policy is to develop listing packages
on individual species. I think the best approach
would be an ecosystem concept where there are two
or three species, such as in Utah, where we have
Thelypodiopsis argillacea, Glaucoc(ir})inn mffrutes-
cens, and Crijtantha barnebyi in the Uinta Basin
that occur in very similar habitats that are close to-
gether. This could be a neat package, and we may
incorporate Cnjptantha grahamii, which is nearby,
and Penstemon gralumiii so you could have four or
five in one package. Now that they have the new
regulations for conducting public hearings, one pub-
lic hearing would take care of all of those.
Q. Ninety percent of the projects are being completed
without an inventory. Isn't that contrary to the law?
A. Not really. It's contrary to in-house policy, but not
to law. The law, of course, is only for listed species.
We have few listed species, and the projects, of
course, are not impacting those. Those are on our
priority list. If they were impacted, we wouldn't al-
low the projects to continue.
Table 2. State programs for T/E plants in the western United States.
States"
Cooperative Agreements
Under Sec. 6 of the ESA
Other State Programs
Animals
Plants
Research
natural
Heritage
areas
Signature stage
Yes, inactive
Signature stage
Active program
Proposal stage
?
Proposal
Idaho Natural Areas
Coord. Committee
Established in 1975,
None
now handled by
state fish and game
None
None
Established in 197.3
State Natural Areas
Preserves Committee
None
Started Fall 1977
Natural Areas
Advisory Preserve
Committee
Established in 1978
■p
Proposal State
p
Arizona
California
Colorado
Idaho
New Mexico
Nevada
Oregon
Utah
Washington
Wyoming
Montana
reviewed
None None
Qualified on 6/2.3/76 None
Qualified on 6/23/76 None
Trying to qualify None
Qualified on 6/23/76 None
None
None
None
None
Trving to qualify Proposal submitted
by state on 5/04/77
Qualified on 6/23/76 Proposal submitted
in 1976
None Proposal submitted
by state on 2/11/75
State program being State program being
reviewed by FWS
°23 states have qualified for cooperative agreement for wildlife programs.
STRATEGIES FOR PRESERVATION OF RARE PLANTS AND ANIMALS
G. Ledyard Stebbins'
.\bstract.- Human preservation of endangered species apparently commenced prior to recorded history with
Ginks hiloba, in China, a tree now known only under cultivation. A niunber of species have become extinct because
man either failed to recognize their value or did not act quickly enough to preserve them even when their value was
appreciated. A philosophy of conservation must be based upon cooperation with others looking to the future. Appro-
priate strategies that could be adapted from the military to achieve the objectives of species conservation include: (1)
Know your enemy, his strengths and weaknesses, and the tactics he is likely to employ. (2) Inferior forces cannot
hope to annihilate or completely neutralize an enemy, but can deflect him from his course. (3) If you have limited
manpower, don't try to do too many things at once; concentrate on primary objectives. (4) Seek the most powerful
allies you can find and learn to cooperate with them as nearly on their own terms as is compatable with your objec-
tives. (5) Soften the enemy by harrassment, when possible, before beginning the final attack. (6) Make use of all the
time that is available; do not risk defeat by premature attack. (7) Never give in as long as there is hope. (8) The most
important principle of all, never underestimate what you are doing.
The first rare species to have been pre-
served by humans was Ginkgo biloba, the
Chinese Maiden Hair Tree. Nobody has ever
seen it as a wild tree. The first Europeans to
see and name it found it in the courtyards of
the temples of China. Fossils indicate that
during the Tertiary period, 30 or more mil-
lion years ago, it was widespread through the
Northern Hemisphere, but by the time hu-
mans had appeared on the scene it was al-
ready confined to China. Where did it grow
as a native and why was it preserved in culti-
vation? A possible answer to these questions
is provided by clues given me by a good sci-
entific friend of mine, the late Edgar Ander-
son. He had one of the most remarkable per-
ceptions for understanding cultivated plants
and their relationships to their wild ances-
tors. He said, "Ledyard, have you ever
thought about this fact— that the trees which
are most successful along the streets of our
cities are those which are native to the banks
of great rivers or deltas? This is because a riv-
er tree is used to being flooded at one season
and parched dry at another season, having
heavy soil dumped on it, and big logs fall
over it, receiving all the punishment that a
tree gets under street conditions." Ginkgo is
such a tree. I was impressed by this many
years ago when I was working at Columbia
University in a laboratory suite on the eighth
floor of the biology building there. I looked
out every morning at the top of the Ginkgo
tree eight stories above the courtyard where
that tree had been planted. In the middle of
New York City, it was certainly a very suc-
cessful tree.
The Chinese plain is traversed by two huge
rivers, the Yangtse and the Hoang Ho (the
Yellow River). Although the climate of China
is a forest climate, those plains are now com-
pletely denuded of native trees. Cultivation
extends right to the edges of the rivers. Pre-
sumably those forests were cut down long be-
fore the Christian Era. My speculation is that
Ginkgo was an element in ancient Chinese ri-
parian forests. When the forests were being
cut down, the priests of the temples thought
it an unusual tree, and having medicinal
properties. They brought in the seeds and
saved trees in the temple courtyards. They
were the first conservationists I can think of.
We come from a long and honorable lineage.
Nevertheless, the concept of conservation
became almost extinct during the earlier cen-
turies of our own millennium. The past 500
years have witnessed the most extensive ex-
tinction of animal species due to a single
cause to have happened during a 500-year
period throughout the evolutionary history of
'Department of Genetics, University of Califomia, Davis, California 95616.
87
88
Great Basin Naturalist Memoirs
No. 3
animals. That cause is human interference
with nature. Among the well-known victims
of human destruction are the dodo, the great
auk, the passenger pigeon, the plains bison,
the moas of New Zealand, the Asiatic lion,
the European forest horse, and the native
wild horses that formerly roamed the Ameri-
can plains. Many others could be mentioned.
Must this destruction continue or can we stop
it? Before considering means of reversing this
trend, we must be fully aware of the problem
we face. Our opponents are not merely a few
greedy men who are out to make a fast buck
in total disregard of other human values. We
certainly must face and neutralize such
enemies by any means available to us. In ad-
dition, we must realize that our efforts are
running counter to a life-style that was
adopted by prehumans long before our own
species. Homo sapiens, came into existence.
Recently acquired knowledge about human
evolution suggests strongly the belief that
when human ancestors left the shelter of the
tropical forests that were their original
homes and began to live by hunting game in
savanna areas, they adopted, partly in a sub-
conscious way, a life-style that was based
upon two objectives: destroy or annihilate
the enemy and exploit the resource. The first
enemies of humanity were predators. Hence
the extinction of the Asiatic lion a short time
after the Christian Era and in very recent
times, the destruction of the California griz-
zly bear, the symbol of our state, during the
19th century.
Many more species have become extinct as
a result, either directly or indirectly, of the
philosophy: exploit the resource. Before hu-
mans started to cultivate fields or domes-
ticate animals, the resources were wild game
animals as well as wild plants that provided
edible seeds, fruits, and roots— consequently
early exploitation of wild horses took place in
most of their range. Anthropologists have un-
covered, particularly in the new world, many
sites suggesting that primitive humans drove
horses over cliffs, slaughtering them whole-
sale, picking up the bodies of those they
could carry and using them for food, hides,
and other purposes. This to them was a nor-
mal way of life. The destruction of flightless,
slow-moving island birds such as the dodo
and the great auk by European sailors and
the moas in New Zealand by colonizing
Maori people came about as a natural result
of a desire for fresh meat on the part of men
who had been deprived of it for a very long
time. Other species have become totally ex-
tinct or preserved only in cultivation or do-
mestication because their existence involved
competition with resources of agriculture or
domestication. Ginkgo has already been men-
tioned. Another is probably the ancestor of
domestic cattle. As is shown so beautifully in
paintings made by men who lived in Europe
from 15,000 to 20,000 years ago, wild bulls
were hunted as game. Archaeological records
suggest that the first domestication of cattle
was connected with religious rites. Bearing
curved horns that resembled a crescent
moon, cattle were regarded by some ancient
tribes as sacred to the moon goddess. Sacred
bulls of ancient Crete are well known to his-
tory, and, in India, sacred cows that cannot
be killed still cause trouble. The expression
"holy cow" is more than a casual bit of mod-
ern slang. It follows a long and venerable his-
tory. Domestication of cattle was one cause
for the near extinction of wild animals. A
beast as strong as a bull could be handled
only if reasonably tame, yet whenever cows
belonging to herds of domestic cattle were
covered by wild bulls and produced calves
from those bulls, genes for wildness in-
troduced in this fashion must have counter-
acted the effects of primitive husbandmen to
breed tractable herds. Wild cattle became
not only a resource to be exploited, but also
enemies to domestication. Modern history
gives us similar examples of species that have
become extinct or nearly so because of com-
petition with various kinds of human efforts.
The senseless slaughter of plains bison and
passenger pigeons during the last century
were not the only cause for the extinction of
these species. Nesting grounds for the passen-
ger pigeon were in rich bottom lands highly
suitable for agriculture. Farm produce was
regarded not only by the farmers themselves,
but by everybody as more important than
nests of pigeons. The buffalo competed with
both cattle and dry farmers. Once the
prairies were fenced in, the wanderers could
no longer survive because their basic way of
life had become impossible. More recently,
plant species have become extinct because
1979
The Endangered Species: A Symposi
89
growing cities have destroyed their habitat.
Of two species that once grew only within
the crowded city hniits of San Francisco, one
of them, Sanicuki maritima, is completely ex-
tinct and another, Arctostapliylas francis-
cana, the San Fransisco Manzanita, is repre-
sented by a single wild shrub plus many in
gardens.
These examples should teach us the follow-
ing lesson: the main barrier to preserving our
priceless heritage of rare animals and plants
is not human greed. It is, rather, the natural
tendency for people of all kinds to be short-
sighted and to prefer to satisfy immediate
needs rather than long-term benefits, particu-
larly those which will be enjoyed only by
their progeny of successors.
Moreover, conservationists are allied with
a whole series of people who realize the need
for reversing a life-style that has dominated
humanity for over a million years. Annihila-
tion and liquidation must be replaced by col-
laboration, or at least tolerance on all fronts.
Exploitation must be replaced by con-
servation. The future existence of humanity
depends on the success of efforts toward this
reversal. Saving rare plants and animals is a
small but highly significant part of mankind's
vital efforts to survive during the coming
centuries.
From the above considerations, a philoso-
phy of conservation must be based more
upon cooperation with others and looking to-
ward the future. Education that might con-
vert apathy into a true realization of the
problem is preferable to a direct attack on an
enemy, who is painted in black colors of un-
compromising greed. Conservation Is a form
of politics, whether we like it or not. In a de-
mocracy or in a community of free nations,
political action, especially when it is prac-
ticed of necessity by a small minority, de-
pends for its success on adopting and exploit-
ing to the limit strategies that are
appropriate for each particular goal. The fol-
lowing well-known military strategies are
particularly appropriate for conservationists
who are seeking to preserve rare animals and
plants. First, know your enemy, his strengths
and weaknesses, and the tactics he is likely to
employ. Potential enemies are any group of
people who for reasons that may seem to be
completely valid and justifiable are likely to
destroy rare plants and animals and their nat-
ural habitat. Among them are people en-
gaged in agriculture, livestock raising, oper-
ators of mines or quarries, prospectors, urban
and suburban developers, developers of mass
recreational facilities such as golf courses and
ski slopes, and conservationists who believe
the greatest need for future civilizations is
water backed up in giant dams and water
power projects. Each of these groups is
armed with verbal weapons that may appear,
on the surface, to be equally or more pow-
erful than any of those in our arsenal.
The growing population needs more food.
New resources of minerals and energy pro-
vided by coal, oil, and gas are vital to the na-
tion's growing economy. The greatest need of
the United States is more and better housing.
Recreational development such as golf
courses and ski slopes make life more enjoy-
able for millions. The rare plants and rare an-
imals of the wilderness can be appreciated
and enjoyed only by a small cult of nature
lovers. The realistic way to provide for future
generations is to build dams that will make
more water and power available to people.
All of these arguments sound logical, realis-
tic, and incontrovertible. In a way they are.
Attempts to refute them directly will cer-
tainly end in failure. Our strategy must be to
recognize the partial validity of these and
similar arguments. We must work around
them, not try to overthrow them.
Tlie second principle is: inferior forces
cannot hope to annihilate or completely neu-
tralize an enemy, but they can deflect the
enemy from its course. On this basis, reason-
able answers to the arguments mentioned
above could be found. Surely more food,
minerals, energy resources, and housing are
needed, but with few exceptions these can be
had by developments that do not destroy pre-
cious and irreplacable habitats and the native
species they contain. The same can be said
even more justifiably about the development
of golf courses and ski slopes. As a matter of
fact, I have taken part in opposition to devel-
opment projects that are ill advised and ill
conceived from a strictly economical point of
view, regardless of conservation. Opposition
from conservationists in those cases called at-
tention to the unsound nature of these proj-
ects and, by causing them to be abandoned,
90
Great Basin Naturalist Memoirs
No. 3
saved the developers or their innocent cHents
from economic embarrassment or possible
disaster.
One of these is located in the north coast
ranges of California. It was a proposed resort
development on the shore of Boggs Lake, a
large, vernal pool. In April or May, during
the wet season, Boggs Lake is a sheet of blue,
limpid water almost a mile in diameter sur-
rounded by gently sloping gravelly beaches,
behind which is a cool pine forest. Situated in
the mountains almost 3000 feet above level,
far from the nearest city or freeway, it would
seem to be an ideal place for a hideaway
where a cool forest glade and pure mountain
water could be enjoyed. That description,
and accompanying photos used by developers
were based only on its spring condition. Its
appearance, however, is highly deceptive.
Boggs has no spring-fed inlet and is exposed
continuously for five months throughout the
California summer to a hot, dry sun. If one,
therefore, visits this "lake" in August or Sep-
tember, the former lake has become a dry
and dusty flat with a few soggy places in its
center. Pine forests are still there, but they
too have become hot and dry and present a
continuous fire hazard.
Pools of this kind usually harbor several
rare and endemic species. Boggs Lake is one
of the best of this kind for botanical research.
When members of the California Native
Plant Society heard about a proposed resort
planned along its shores, we went in great
number to a hearing in the Lake County
Courthouse to present our views. Before
doing so, we took the trouble to walk around
the area carefully and acquired a greater fa-
miliarity with the terrain than had the devel-
opers. Their publicity was based chiefly upon
an airplane survey. Our view of the situation
was strong enough to dissuade both the coun-
ty supervisors and the developers from con-
tinuing the project. Boggs Lake was then ac-
quired by the Nature Conservancy and its
imique habitat is premanently preserved.
The third principle is, if you have limited
manpower, don't try to do too many things at
once. Concentrate on primary objectives. In
terms of conservation strategy, do not spend
valuable time on every species that is rare
and local. Most of the rare species that live in
national parks, state parks, wilderness areas,
that have been set aside by the national park
and similar privately controlled areas, need
only occasional monitoring to see that provi-
sions and rules for preservation are being car-
ried out. Sometimes the officials need to be
informed. I remember an example of a grass
in the Sierra Nevada, a rare species, Stipo la-
tiglumis, known from only about three local-
ities. I had a suspicion about its origin. I sus-
pected that it evolved in what now is a
genetically familiar fashion: crossing between
two other species of Stipa and doubling the
chromosome number: an allopolyploid. The
most accessible place for this species, accord-
ing to herbarium labels, was Lost Arrow
Camp in Yosemite Valley. In Yosemite Na-
tional Park, as in other parks, a collecting
permit is required. Collecting permits always
say in very large capitals, NO COLLEC-
TING OF ANY KIND IS PERMITTED ON
THE FLOOR OF THE VALLEY. Never-
theless, I went to the park naturalist's office
to ask for a permit. When I explained what I
wanted, the park naturalist himself received
me. He said, "Where does it grow in the val-
ley?"
"The labels say Lost Arrow Campground."
"This is where the government center is
built."
"Do you think there are any native areas
here?"
"I think I know them pretty well, but I
don't think you'll find anything unusual here
at all."
"May I look? And if I find it here, may I
collect'it?"
"Well, I guess you can."
We started looking. We found it in the
front yard of the private residence of the
park naturalist himself. Its allopolyploid na-
ture was demonstrated by Dr. Richard Pohl.
Other rare species, not in the national
parks or preserved areas, nevertheless grow
in such inaccessible spots that they are very
unlikely to be destroyed. An example is a
species of the genus Eupotorium that many
years ago I discovered on a north-facing
limestone cliff near Lake Shasta. Eupatorium
shastensis is always perched on cliffs, and 80
percent of the plants of it are so high up on
the cliffs that no one can reach them except
by specialized rock climbing techniques.
There is danger, possibly, from prospecting
1979
The Endangered Species: A Symposium
91
or blasting of these limestone cliffs, except
for the fact that they are in very rugged ter-
rain, one of them isolated from any highway
by the waters of Lake Shasta and the other
on the summit of a very rugged mountain.
Bringing in equipment to mine these areas
would be extremely expensive. Because it is
on their land, the Forest Service knows about
it and I believe will not issue permits for
prospectors or mining on these rather unusu-
al limestone cliffs. This case requires mon-
itoring, even if there is no formal preserva-
tion.
General applications of this strategy, I be-
lieve, is to keep lists of rare and endangered
species as short as practicable, to pay as
much attention as possible to the amount of
danger and the nature of the danger to which
a species might be exposed and to determine
actual rarity in terms of space occupied and
actual numbers of individuals in each popu-
lation. Government officials and leaders of
general conservationist organizations, such as
the Sierra Club, should not be presented with
lists of two or three hundred species with un-
familiar names. I suspect that in many in-
stances these are filed in some cabinet, which
a secretary might open every six months or
so. Here is a situation where the more we
know about potential and imminent danger,
the better off we are.
The fourth principle is to seek the most
powerful allies you can find and learn to co-
operate with them on as nearly their own
terms as is compatible with your objectives.
My happiest experience with powerful allies
resulted in partial preservation in an area
that for 25 years previously had been very
dear to my heart. This is a little-known por-
tion of the fabulously scenic Monterey Penin-
sula on the coast of central California. That
area, a small, ancient "raised beach" millions
of years old (Pliocene), is underlaid by a ster-
ile, hard, and impervious "hard pan" soil. Its
plant communities contain so many problems
in evolution and plant geography that I have
nicknamed it "Evolution Hill." Its most dis-
tinctive tree species are the Bishop pine and
the narrowly endemic, rare dwarf, Gowen
cypress. Each time I have taken students to
this area it has given me cause for apprehen-
sion. We could traverse by foot a network of
trails and rough roads. The owners had the
trees and bnish cut so that they could very
easily be converted into paved streets and the
whole place put into a resort development.
Ownership is in the hands of an exclusive
multimillion dollar organization, Del Monte
Properties, which was then the fiefdom of
one of the most prominent citizens of north-
ern California, Samuel F. B. Morse. One day
during the 1950s I obtained an appointment
with Mr. Morse to discuss the future of Evo-
lution Hill. The great man was polite and
cordial. He said that he too was much inter-
ested in saving the area and to see that it re-
mained preser^/ed as long as he remained in
control. He could not, however, make com-
mitments that would tie the hands of his suc-
cessors. Mr. Morse at that time was in his late
seventies and he had clearly given me only a
temporary stay of execution. Several years
later, after Mr. Morse's death, the blow fell. I
received a telephone call from a prominent
resident of the peninsula, the director of a
nearby laboratory. He said, "I want you to
come down to Salinas to attend a meeting of
the County Planning Commission. The new
director of Del Monte, who used to be vice-
president of the Corning Glass Works, wants
to start a sand quarry for glass in the forest
right behind our house." I realized at once
that Evolution Hill was in danger, but also
that we members of the California Native
Plant Society had powerful allies. Several of
the most wealthy and prominent home-
owners who had bought and built in the for-
est in order to have quiet solitude with undis-
turbed woodlands for hiking and horseback
riding felt that their life-style was severely
threatened and that the hundreds of thou-
sands of dollars they had invested in their
homes might go down the drain. The result
of the first hearing was noncommital, but the
stay of execution was maintained. No permit
to quarry was issued. We then organized a
joint fact-finding site visit attended by more
than 100 members of the Native Plant So-
ciety plus several homeowners. Such an event
deserved and received good newspaper pub-
licity in the area. Hearings and litigation con-
tinued for about two years. Finally the
quarry-minded individuals from the Del
Monte Company gave up the sponge. They
donated a portion of the hill to the county to
be set aside as the S.F.B. Morse Preserve and
92
Great Basin Naturalist Memoirs
No. 3
agreed not to quarry for sand in an area near
the established homes. We would have pre-
ferred to see the preservation of all of Evolu-
tion Hill, and this may still be possible. Ten
years after this partial victory, it is still as I
first saw it; no homes have been built in the
area.
The fifth principle of strategy: if possible,
soften the enemy by harrassment before be-
ginning the final attack. This principle is well
illustrated by an experience we had a few
years ago in an endemic area in the Sierra
Foothills, known as Pine Hill. This hill, 25
miles east of the state capitol of Sacramento,
about 2000 feet high, has a number of en-
demic species. The most spectacular of these
is a flannel bush, Fremontodendron deciim-
bens, described by Dr. Robert Lloyd. It is
noted for its prostrate habit and its copper-
colored flowers, where most flannel bushes
have bright yellow flowers. It is a very dis-
tinctive species, not known anywhere except
on Pine Hill. I say with some confidence that
my friends and I have combed over every hill
in the neighborhood that could possibly hold
it and we have never found it, so I'm certain
the central ridge of Pine Hill is the only
place where this shrub grows.
One day a member of the Native Plant So-
ciety visited Pine Hill only to find that the
Forest Service, in order to construct a fire
break, had cut down almost all of the shrub
of Fremontodendron, and it looked as if it was
gone. His reaction was immediate and posi-
tive. He wrote a strongly worded article that
was soon published in our society's journal.
The article brought a flood of letters from
outraged members of the Native Plant So-
ciety to the office desk of the district man-
ager. That was in May. In October I got a
letter from a friend in the nearest town, Pla-
cerville: "Ledyard, I want you to come. I've
got to go out with the ranger to Pine Hill."
Why? "Because they want to put in a little
powerplant, about 10 X 20 feet and they
want to do it without having all the flack
that we gave them on the fire break." So we
went up there and we told them where to
put it, a place where there were almost no
plants. Soon after, we were able to enlist the
powerful ally. The husband of the secretary
of our Sacramento chapter of the Native
Plant Society, Warner Marsh, had been in
the Sacramento office of the State Forest Ser-
vice for many years and was highly respected
by all personnel in that service. So, Warner
went out with one or two other people and
the ranger and put a little pink ribbon on
every shrub of the Fremontodendron. Fortu-
nately, it is quite a resilient shrub. Cutting
down the branches didn't destroy the roots,
and so new branches came up. They're back
again and now the California State Forest
Service isn't going to disturb them. We are
having other problems with Pine Hill be-
cause of changes in the state government or-
ganization, but we're still very optimistic
that the whole area will be preserved.
The sixth principle of strategy is to make
use of all the time that is available. Do not
risk defeat by premature attack. Many con-
servationists who are aware that an unusual
habitat is threatened by mining, quarrying,
development, or some other way, tend to
magnify the threat and particularly its imme-
diacy. Sometimes this attitude is justified and
necessary; other times it is not. Surely, if the
developer is known to have his eyes on one
of our favorite spots, we must act quickly
with all resources at our command. Never-
theless, we cannot be stampeded by a poten-
tial danger which may not be realized for
some time. Here again precise knowledge of
the danger that threatens a rare species or
community is of the utmost importance.
The seventh principle is never give in as
long as there is hope. One can lose several
battles but still win the campaign. The last
two principles are well illustrated by the
campaign to save the lone Manzanita area on
the eastern margin of California's central val-
ley, one of the most dramatic of California's
ecological islands. I call it an ecological is-
land because the soil is so different from the
surrounding soils that the species living there
are isolated as if they were on an oceanic is-
land surrounded by a sea of grass and oaks.
Another inhabitant of the barrens is a species
of buckwheat, Eriogenum opricum, described
about 25 years ago by J. T. Howell.
When the California Native Plant Society
was formed, one of our objectives I thought
of almost immediately was saving lone Man-
zanita, so a group of us went to the Amador
County Courthouse first to find out who
owned it. The results were not encouraging.
1979
The Endangered Species: A Symposium
93
The whole area belongs to a syndicate con-
trolled by a large San Francisco bank, which
leases land to miners and quarriers because
there is a clay of extremely high value. When
we approached one of the officials of this
company, we got a very emphatic reply,
"We'll mine every blank blank cubic foot of
that sand and clay and we dare you blank
blank s. o. b.'s to stop us."
Somewhat later, we were still trying to
find a way around them and went on a Sun-
day when we thought nobody would be there
to look for another spot for the Eriogonum
epricum. We ran into some people who
turned out to be miners who were not mining
on Sunday, but were hunting quail or some-
thing like that. They said, "What are you
doing in our place?" We explained what we
were doing. "You better get off. We're hon-
est miners and we've been working this for
20 years. This place is full of rattlesnakes and
I wish there were twice as many of them to
keep you blank blank blanks from going on
to it." Well we haven't given up. We've had
articles in our journals. We've had publicity
wherever we could find it. We've discussed it
with the California Department of Parks and
Recreation and other groups, and we have
gained some allies. Meanwhile the quarries
that existed for some time are still there and
still working, but they haven't invaded any
more territory than they had when we first
started in 1966. So while there's life there is
hope.
(NOTE: As this article was going to press,
I received a welcome announcement: The
central heart of the lone Manzanita area has
been purchased by the Nature Conservancy.)
The eighth and last principle— the most im-
portant principle of all— never underrate the
importance of what you are doing. Human
civilization is built on two great pillars. A
pillar of knowledge and a pillar of beauty
and its aesthetic appreciation, whether it be
the beauty of nature, artistic creation, or the
beauty of the spirit. Drs. Lovejoy and Cle-
ment this morning showed us part of a world-
wide effort to save humanity from its own
destruction. Fountains of knowledge can be
bound up in the most ugly and unattractive
weeds we are trying to save.
A plant known only in a few suburban
areas, which is now severely threatened, is a
tar weed known as Holocarphra macradenia.
Now tar weeds are among the nastiest weeds
in California pastures. To try and tell a ran-
cher that you want to save a tar weed is just
like telling him to stop drinking beer. Well, it
so happens that this species was part of a
large-scale research project carried on by I.
Clausen and D. D. Keck 25 or 30 years ago.
They discovered that what the taxonomists
had called two species are actually four mor-
phologically recognizable ones. Among those
four species, hybrids between almost any col-
lections from two different localities were
sterile or couldn't be made. In other words,
hiding under first two and then four species is
a whole series of little narrow endemic spe-
cies, the nature of which is associated with
chromosomal difference. In our quest for un-
derstanding the mechanisms of the origin of
species, the tarweeds, including Holocarpha
macradenia, could be a key group. Now we
will have to resist the desire to succumb to
the developers and keep the species alive, at
least under cultivation. After all, the habitat
will be gone anyway. The place where it has
been known for the last 50 years is in associ-
ation with wild oats and other introduced
species. Its prehuman habitat was gone long
ago. This is an example of a humble sticky,
smelly, nasty weed which could be a gold
mine of scientific information.
Now we should then come to the aesthetic
value. My illustrations cannot equal the beau-
ty you saw in the booklet of the National
Wildlife Federation we all received this
morning. I'll show finally just two slides
which give a modest impression of the beauty
of plant species. One is a Monterey cypress,
growing on the granite cliffs facing the blue
Pacific Ocean, with its picturesque branches
and trunk growing out of solid granite. The
other is a pure white flower of the California
rose mallow centered with the deep maroon
spot in the middle of the flower, growing in
the hot valley in the middle of the summer.
STRATEGIES FOR PRESERVATION OF RARE PLANTS
Arthur H. Holmgren'
.\bstract.— Preservation of the habitat is the only logical strategy to save endangered species from earlv extinc-
tion. Ecological amplitudes of rare species are very narrow, so transplantation to such alien sites as botanical gardens
is not a solution. Protection may not be the answer. We must learn as much as we can about the biology of the
species in question, in the field and under laboratory conditions. The first steps must be to determine the distribu-
tion. This would be followed by analysis of soils by means of physical and chemical studies. Pollination ecology,
associated species, phenological records, and genetic and cytological studies must be a part of the biological studies.
Such studies would require teamwork by qualified botanists.
I suspect I was asked to take this assign-
ment because, as several of you know, I have
cultivated many of our western native plant
species. Most of these plants were introduced
into my gardens so I could have laboratory
material for my taxonomy classes. I had great
success with Penstemons and at one time I
had 33 species in this genus. Many of my
Penstemons hybridized under prolonged
flowering conditions in my gardens until it
was difficult or nearly impossible to deter-
mine parents of most of my hybrids. Some of
my introductions in other genera became
troublesome weeds. These are not the kinds
of species we are concerned with in this sym-
posium.
I will devote my time to strategies for
preservation of rare plants. My answer and
only logical strategy is to preserve the habi-
tat of the threatened and endangered species
so that we may save them from early extinc-
tion. Species inevitably become extinct, in
times past by natural forces, but in recent
times greatly accelerated by man's destruc-
tive activities.
Extant knowledge of rare species indicates
that ecological amplitudes are very narrow
and thus transplantation to such alien sites as
botanical gardens is not a solution. And still,
Franklinia alatamaha Marsh was preserved
in cultivation. The lost camellia or Franklin
tree, originally from someplace in the coastal
plain of Georgia, was discovered by John and
William Bartram in 1765 and has not been
seen in its native place since 1790. Many bot-
anists have searched long and hard for the
lost camellia. Dr. Ritchie Bell of the Botany
Department at the University of North Caro-
lina has made several expeditions with gradu-
ate students in search for the lost camellia
that has been in cultivation for nearly 200
years.
In the absence of hard data, habitat preser-
vation is the only option open, and it is in-
creasingly at hazard because not even the sci-
entific community understands the problems.
Habitat preservation is seen as a powerful
threatening tool to the. public at large and es-
pecially to those who are anxious to develop
our natural resources. Elected office holders
and seekers are afraid to line up with the bi-
ologist who sees the need to preserve habitats
of threatened and endangered species. We
have no idea yet how much area to protect
or even if protection is the answer.
Two species come to mind that thrive in
disturbed sites. Astragalus patjsonii (Rydb.)
Barneby is usually found in burned-over areas
in Wyoming and Mertensia toijahensis
Macbr. thrives in similar habitats in the
Toiyabe Range in Lander County, Nevada.
Many species make a living in disturbed sites,
but it is unusual to find rare species in .such
habitats. Perhaps more fieldwork will show
that the two species mentioned here are not
as rare as we have thought.
Dr. Howard S. Irwin, president of the New
York Botanical Garden, said in a letter to me
'Professor Emeritus of Biology and Acting Curator of the Intermountain Herbarium, Utah Stale University. Logan. Utah 84322.
95
96
Great Basin Naturalist Memoirs
No. 3
dated 16 October 1978: "The most desperate
need is a federal program that would encour-
age students to study the biology of species'
rarity, more or less in the manner followed
by Dr. Lazarus Walter Macior at Akron Uni-
versity in investigating the Furbish louse-
wort."
I contacted Dr. Macior and received a
prompt reply dated 14 November 1978. Dr.
Macior enclosed a copy of his manuscript
which will appear in the October-December
1978 issue of the Bulletin of the Torreij Bot-
anical Club. All papers published in the Bul-
letin of the Torrey Botanical Club are copy-
righted, so the manuscript was sent to me
for my personal use only. Dr. Macior's work
with the Furbish lousewort will certainly be-
come a model for experimental studies on
rare species.
The first step in a strategy for preserving
rare plants must be to learn as much as pos-
sible about them in the field and under con-
trolled conditions in the laboratory. Every ef-
fort must be made to determine the
distribution of the taxon in question. Sme-
lowskia hohngrenii Rollins was thought to be
confined to one rock prominence in the To-
quima Range, but Sherel Goodrich discov-
ered that this unusual species was actually
more common in the Toiyabe Range to the
west. The known distribution at this writing
includes four stations in the Toquima Range
and 10 populations in the Toiyabe Range, so
the species is not considered to be in the pre-
carious situation suspected prior to the 1978
field season. This unusual mustard is a dis-
tinctive species, and, as is true so many times
in this family, species are easier to recognize
than the problem of assigning them to gen-
era. I still have difficulty thinking of this spe-
cies as belonging to the genus Smelowskia.
To me, it seems to have closer affinities with
the genus Bray a, far to the north. It may turn
out that we have a new genus. Arahis
shockleyi Munz is another species that may
turn out to be more common than we have
thought. Only a few widely scattered collec-
tions have been made from Tooele County,
Utah, to the San Bernardino Mountains in
California. The paucity of collections prob-
ably illustrates how poorly some of our desert
ranges are known.
Detailed field studies would vary to some
extent with different species. The Smelowskia
of central Nevada is found in crevices of an-
desite rocks, and future studies on this species
may show that it has a preference for this
kind of a substrate. Arctomecon humilis Cov-
ille and A. californica Torrey and Fremont
probably require gypsum soils, often referred
to as "gumbo" clay.
Detailed biological studies would begin af-
ter the distribution of the species has been
determined. Biological studies of rare species
would investigate the ecological adaptations
of the species as to edaphic factors and biotic
environment. Soil samples would be taken
from many sites and thoroughly analyzed
with every sophisticated chemical and phys-
ical means we know. Weather records would
be analyzed or gathered. Total precipitation
means little unless we know the distribution
throughout the year. Climatic characteristics
in a broad sense would also include solar
radiation and temperature records.
Pollination ecology may be a key as to
why a species is rare and perhaps even on the
verge of becoming extinct. The loss of a pol-
linator through spray programs may place a
species in imminent danger of becoming ex-
tinct.
Phenological records should be kept and
associated species recorded. What are the re-
quirements for seed germination? Much re-
mains to be learned about seed germination
and especially for rare species. Under what
conditions is flowering initiated? How soon
after flowering are fruits matured, and what
is the mode of seed dissemination? Are cer-
tain species usually associated with a taxon
we are studying, or is a niche so inhospitable
that our species has the habitat without a
competitor?
Cytological studies would help in possibly
determining closely related species. Dr.
James Reveal and I prepared a paper several
years ago on Cilia caespitosa A. Gray that
was never submitted for publication. The
chromosome number of this rare and restrict-
ed species was determined to be the same as
the wide-ranging and highly variable G. sub-
nuda A. Gray, 2n = 16. Gilia caespitosa is
restricted to white, decomposed sandstone
one mile south of Teasdale in Wayne County,
Utah. Dr. Dieter H. Wilken has prepared a
fine paper on G. caespitosa in much more de-
1979
The Endangered Species: A Symposi
97
tail, "The Status of Gilia caespitosa A. Gray
(Polemoniaceae)," which has been accepted
for pubHcation in Madrono. Dr. Wilken has
concluded that G. caespitosa has a close rela-
tionship with G. subnuda. Reduced speci-
mens of G. subnuda are very similar to the
uniform specimens of G. caespitosa. Dr. Wil-
ken suggests that G. caespitosa may represent
one of he more primitive elements within
Gilia. In my Honor Lecture in 1977 at Utah
State University in Logan, Utah, I pointed
out that this beautiful little perennial may
have arisen from an extreme biotype of the
variable and common G. subnuda. If a wide-
spread species becomes established in an un-
usual edaphic situation, it will carry only a
small part of the genetic variability of the
original species. Inbreeding and random fix-
ation will tend further to make this insular
population more uniform and still more dif-
ferent from the ancestors as the years of iso-
lation continue. The individual plants of this
beautiful species appear to be as nearly gen-
etically alike as separated parts of a clone. It
does not seem logical that this species is a se-
nescent species. It probably evolved where it
is found today and adapted to a single eco-
logical niche. It seems unlikely that it occu-
pied a larger area in past times.
The biological studies outlined above are
all a part of the first phase in learning about
rare species, but the studies obviously do not
stop here. If the species is threatened in part
of its range, some natural populations must
be preserved in situ for further study. This is
especially true for the three species of Ar-
ctomecon. Each species is confined to a pre-
carious habitat and all three are fast dis-
appearing. The fact that requirements for
seed germination are not known and that the
plants cannot be transplanted make it im-
perative that these plants be studied in situ.
Housing developments, trail bikes, and gyp-
sum processing plants doom at least two of
these bear poppies. Oh, yes, I had better not
forget to point a finger at the plant tax-
onomist. I was appalled when I discovered
how many specimens we had in the Inter-
moutain Herbarium. It is well known that
numbers of specimens in a herbarium are no
indication of rarity.
Another step in preserving a rare species
may be in attempting to cultivate plants in
identical habitats or very similar to the origi-
nal ones. This would involve attempts to ger-
minate seeds for transplanting of garden or
greenhouse-grown plants and transplanting
growing plants from natural habitats. This
does not excite me, as we have attempted to
grow several rare species. Leila M. Shultz,
curator of the Intermountain Herbarium, suc-
ceeded in germinating seeds of the rare
Sphaeromeria mtJiiae Holmgren, Shultz, and
Lowrey from Zion National Park. At the
present time, we have two potted plants
growing in my home greenhouse in soil from
the type locality. After two years of vacil-
lating from "Looking good" to "Will they
make it?" I begin to wonder if my specimens
will ever flower. So many things come to
mind. What about solar radiation, length of
day, and on and on? I have successfully trans-
planted and multiplied Cijpripedium cal-
ceolus L. from the mouth of the Logan Can-
yon that was in the way of a new home. I
have divided the clumps several times and
even moved the entire population when we
moved from our old Logan home to a site
near the base of the mountain just north of
the mouth of Logan Canyon. There are more
plants today in my garden than the original
population contained 35 years ago. I have
thought of moving a few plants to sites in Lo-
gan Canyon to habitats that would probably
support this lovely orchid, but I hesitate
when I think of the pressure of every foot of
bank area along Logan River by fishermen.
The plants I am growing represent the only
known living plants of this species in Utah.
Extirpation would once again remove a spe-
cies from the wild that ranged from Logan to
Provo when the Mormon pioneers came to
Utah.
I have attempted to grow the rare hetero-
stylous Primida maouirei Williams that is
known only from a nine-mile stretch in Lo-
gan Canyon and consisting of only seven
known populations. Plants flowered the first
year and emerged the second year without
flowering, and, after languishing for a short
time, disappeared from my garden spot,
which I had thought was quite similar to the
canyon habitats. 1 hope to see a graduate stu-
dent work out the biology of Primula ma-
guirei in the near future. Howard Irwin re-
ports that the New York Botanical Garden
98
Great Basin Naturalist Memoirs
No. 3
recently got a grant to conduct studies of the
New York Monkshood, Aconititm novabores-
cense A. Gray, initially to determine its dis-
tribution and also to get some biological
study imderway. This unusual monkshood is
presently known from a few localities. This is
the way to go, and sometime in the future we
will have hard data to give us a better under-
standing of past and present histories of floras
and species.
The most important strategy of all has to
be for us to win public support and thereby
gain support of elected officials on all levels.
Unless we gain this support, there will be no
funding for the work that is just beginning.
We have made gains since Earth Day, 22
April 1970, but in other important ways, we
have lost ground (no pun intended here). The
radical rhetoric of street protests has been re-
placed by legal briefs. There are probably
more than 8 million members of environmen-
tal groups who make contributions totaling
nearly 70 million dollars a year. The Audu-
bon Society and Sierra Club were the first of
the conservation groups, but they have been
joined by many more. State native plant so-
cieties are organizing, with several new so-
cieties each year. We know of the accom-
lishments in California and what the
potentialities are. We have just organized a
Utah Native Plant Society. This is the way
we can get our message to the state and local
levels.
I share the anxiety of Howard Irwin, Walt
Macior, and Dieter Wilken in preserving rare
species; but, in the meantime, we cannot be
sympathetic with those who would preserve
them only as instruments for political strate-
gy. Those who have taken strong stands pro
or con without sufficient knowledge have
hurt our cause. In the meantime, let's study
our rare species intently with qualified, pro-
fessional botanists.
Questions to Dr. Holmgren
Q. There is a big problem in that information available
is not keeping up with the demand. The gentleman
from the Forest Service said they had 200 cited to
survey and funds to do 20. The problem is even
greater in private industry. The company proposes a
project and requires a survey and the information is
just not generally available. Do you foresee a way
out of that dilemma?
A. I just don't see a way out of it. In fact, very often we
see proposals or requests for proposals come across
our desks and we are supposed to have something in
on it a week before the proposals came to us. Some-
times we have about six weeks to work this out.
There is no way we can do it. To pretend that some
of these things can be done in such a hurry is not
being honest with the problems that are at hand. It
is going to take some time. Very often these things
have been under planning stages for a long, long
time, but the problems do not come to us until the
last minute. No one plans a $100 million plant with-
out having gone through a lot of planning, and then
in the final stages the requests come across our
desks. What can we do? It is going to take some
time.
Comment: The Forest Service is developing a policy
now that would require all external organizations
proposing projects on Forest Service land to hire a
professional botanist to inspect the project for T/E
species, so we'll get a lot of these covered in that
way.
Q. Does the cultivation of plants and plant planning
hold a better opportunity than we have experienced
with animals?
A. Sometimes it does. Janice Beetley brought in some
seeds oi Arctomecon, and she succeeded in germinat-
ing them but they never flowered for her. We know
that is a genus where transplanting is an impossibi-
lity. It surprises me because so many members of the
poppy family can be grown from cuttings, but this
particular one defies that. I used to think I could
grow anything if I knew the right witchcraft, but
I've discovered there are all degrees of absolute suc-
cess, to the point where you have weeds coming
along in your garden to the point of absolute failure
on the other end. In my years of experience with na-
tive plants, I could plug in something all along the
wav so that I would go imperceptibly from complete
success to failure.
Comment: A comment really to the gentleman's earlier
comment. I believe there are a growing number of
industrial concerns who recognize the problem of
endangered species to the point that they would
much rather incorporate biological knowledge ear-
lier in the planning process than face litigation later
on. In this way I think there is progress being made
in this area.
A. I'm sure there is. I think that is one of the good
things about some of the problems we've had along
the way that these people have discovered. As they
begin to plan, this is one part that has to be in the
planning stages right from the very beginning. I
think there were references to that in talks we heard
yesterday. People are beginning to come to some of
these agencies, and Doug Day has had several
people come to him and ask for help as they were
beginning to plan a study. I think we are going to
have more of that to a point where I hope that final-
ly we can get the public on our side. It's going to be
a long education, but every day when I pick up
newspapers now I read articles by different authors,
DeLong and several others, who are writing very
well-written essays on the problems we are now fac-
ing.
1979
The Endangered Species: A Symposium
99
Q. One thing I'm surprised no one has mentioned. I'd
hke to know if the Fish and WildUfe Service has
contacted either BYU or the Intermountain Herba-
rium. \t the present time Stan Welsh has com-
puterized all the herbaria for Colorado, Wyoming,
North Canada, and North Dakota. Listed in their
computer program is every sample surveyed of those
herbaria. Any agency or industry person interested
in developing a project need simply place a phone
call to Fort Collins asking them to print the species
list. It has a tremendous option on it. Included on it
are all the rare and endangered species in a specific
geographic area and I've heard nmiors from work-
shops held in Fort Collins that they plan on expand-
ing this. Have you been contacted about that?
.\. I haven't. Have you, Leila Shultz?
Comment: No, but I do have a comment on it. Herbaria
standardly have 40 to 70 percent misidentifications
and so, as good as the information is, it's nice to
have it available. But if you want lots of mis-
information you can get it quick.
A. Yes, there are so many people who will look at a her-
barium label and the identification on that becomes
the gospel truth. We know. We get plants from oth-
er institutions that are not even in the right genus,
and sometimes the species is a long ways away (but
not from the BYU). We're glad we have such good
working relationships with all universities.
Comment: I have talked to Colorado State about possi-
bly getting on this sytem, and, although there are
misidentifications, I think, where you have the com-
puter printout, if something comes out in the distri-
bution very different from what you expect, it comes
to your attention in a hurry. I think there is good po-
tential in it.
\. I was trying to get Leila's attention because for the
last several years she has been listed as the assistant
curator, but I'll have you all know that she is the
curator. She's done it all and I've been happy, but
sometimes it has given her more than a person ought
to carry.
Comment: I have one comment here. Perhaps the infor-
mation or the lack of information here with regard
to the private industries approach to endangered
and threatened plants needs to be traded. Until now
it hasn't been, so I will take it upon myself to give
you my own professional view of it with regard to
the private industries I have dealt with. Private in-
dustry is willing to cooperate with the endangered
species program. They do not wish to interdict any
of the endangered or threatened species. They are
willing to do what is necessary in order that they
may fall in place, but they do need to be able to sur-
vive the regulations so that they can carry on their
businesses. The problem arises though, not with the
private industries, but with the general public. The
general public is the place where we really need to
do our education job and not with the private in-
dustries. The private industries are ordinarily with
us.
STRATEGIES FOR THE PRESERVATION OF RARE ANIMALS
Clayton M. White'
,\bstract.— Strategies used to enhance or help restore rare and endangered animal species are reviewed. No new
strategies are presented, but rather a review of various levels at which programs can be initiated are indicated.
Nearly 93 percent of the recognized endangered animals are vertebrates. Programs to help restore vertebrates can be
aimed at either the habitat or the organism level. Habitat restoration or preservation is the more difficult to achieve
and, accordingly, most strategies are aimed at the organism. Species, populations, or communities and ecosystems
can be treated, but most often each requires a separate approach. Among organisms the species level attracts most of
the enhancement effort because species are easier to understand and deal with. Numerous strategies are being tried
with species, and several examples, such as the Aleutian Canada goose, Galapagos tortoise, a wingless undescribed
orthropteran, and fish species requiring turbid water, are given. Populations or communities of animals are more
difficult to work with, but some strategies such as faunal reserves are likely to be successful. A table listing .39 exam-
ples of endangered species from across the animal kingdom, along with the major reasons for their declines and
currently working strategies or possible ones to help them recover, is presented.
This report reviews strategies used to en-
hance or help restore rare or endangered ani-
mal species, and, as such, the approaches are
clearly different than most of the strategies
reviewed by Holmgren (this volume) for
plants. This report, however, is not intended
to be an exhaustive review of all the various
plans, mechanisms, or strategies that have
been used with all animal species.
At the outset, it must be recognized that
most ideas advanced thus far apply to verte-
brates and are not necessarily applicable to
invertebrates. In part, our knowledge of the
population dynamics of many rare in-
vertebrates lags behind that of the verte-
brates. As pointed out earlier by Lovejoy
(this volume), the invertebrates are also cur-
rently treated administratively in a different
manner than vertebrates. Those animals offi-
cially recognized as rare or endangered by
federal or international conventiorr as of 1
December 1978 (U.S. Department of the In-
terior 1978) are in the following categories:
lammals
irds
leptiles
Jiiphibians
•ish
281 Snails 8
214 Clams 25
68 Crustaceans 1
16 Insects 8
51
'he above total
to 672 kinds, plus there are
an additional 158 kinds proposed. Of those
listed, about 74 percent are mammals and
birds and nearly 93 percent are vertebrates.
Before approaching the topic of discussion,
I think it fair to conclude that endangered
species programs and concerns are mainly
generated in affluent societies where people
have the leisure time and the monies to con-
sider such problems (cf. Smith 1976). Roland
Clement (this volume), however, ironically
pointed out that it is the affluent societies
themselves which, because of their exploitive
nature, have been the root of the causes of
habitat destruction and thus species endan-
germent. The basic question seems to be
"What will man as a species tolerate?" In
nonaffluent regions they appear to tolerate
considerable, even the loss of part of their
faunal heritage. Myers (1971), in discussing
the preservation of fauna in Uganda, has cor-
rectly pointed out that the wildlife heritage
of that country is of international concern,
and that the land .should be developed, con-
served, and managed in accordance with
sound ecological principles. Sociological con-
cerns also play an important role in such is-
sues.
Another curious paradox is that the bald
eagle {Haliaeetus leucocephalus), which we
have declared as part of our heritage by af-
'Department of Zoology, Brigham Young University, Provo, Utah 84602.
101
102
Great Basin Naturalist Memoirs
No. 3
fording it the status of the symbol of our
country, is endangered. Similarly the quetzal
{Pharomachriis mocinno) is endangered in
Middle America, where it is represented on
the coinage and is a national symbol in
Guatemala (Lovejoy, this volume).
Levels of Organization
There are at least two major levels of or-
ganization from which strategies for preser-
vation have been or can be approached,
namely, (1) the habitat (environment) level
and (2) the organism level.
The only clear way to preserve animals is,
of course, to preserve or maintain habitat in
large enough blocks of land to maintain the
species diversity. This leads basically to the
concept of the "megazoo" as developed by
Sullivan and Shaffer (1975). Among others.
Diamond (1975, 1976), Terborgh (1974), and
Wilson and Willis (1975) also discuss the hab-
itat approach to organism preservation and
the strategies achieved by maintaining signif-
icantly large tracts of habitat.
The habitat and the organism level have
rather different approaches. For example,
zoological gardens may be used if simple
preservation of the rare or endangered ani-
mal is desired (Conway 1967, 1978). Such
preservation, however, can be and most fre-
quently is independent of any approximation
of habitat because the animals are simply
bred and maintained in cages. Nonetheless,
there are currently 26 species of rare mam-
mals that have self-sustaining populations in
zoos (Finder and Barkham 1978). These rep-
resent one species of marsupial, 2 primates, 8
carnivors, 3 perissodactyles, and 12 arti-
odactyles. In some cases, there may be as
many or more individuals in zoos than there
are in the wild. The Asiatic lion has 96 in
captivity and an estimated 177 in the wild,
the Siberian tiger has about 450 in captivity
and perhaps 200 in the wild, and the Pere
David deer is extinct in the wild but about
777 remain in captivitv (Finder and Barkham
1978).
Before strategies for preservation can be
formulated, it is necessary to determine the
reasons for endangerment and that, in turn,
may lead to a basic knowledge of the biology
of the animal to be dealt with specifically. It
is difficult to understand an animal's rareness
when we don't understand its population dy-
namics (see Drury 1974 and Smith 1976).
Simon (1969) discussed some of this when he
described the status of 86 rare mammals in
33 countries; examples follow:
Feru — Vicuiia {Vicagna vicugna): infor-
mation inadequate and conflicting,
field surveys needed.
Brazil — Thin-spined porcupine {Chaet-
omys siibspinosiis): little known
about species beyond fact that
range is restricted, deforestation
modifying habitat, and field stud-
ies needed.
Morocco— Barbary hyaena {Hyaena hyaena
barbara): status precarious, true
position obscure, and studies
needed.
Following Simon's (1969) assessment. Mill-
er Rottman, and Taber (1973) studied the
vicufia in South America and determined that
the present stocks are occupying suboptimal
habitat and that the carrying capacity of the
present available habitat (in terms of pre-
ferred forage etc.) is about 10 times the pres-
ent numbers. The current range is less well
watered than the optimum range. Because
the species need to water daily, the answer to
improving the population status seems to lie
in providing watering localities in areas
where forage is adequate but no water exists.
Aside from habitat improvement, hunting for
fur also needs to be controlled.
It is surprising how little we know about
the biology and evolutionary dynamics of
even the most common species. Both the
house sparrow (Passer domesticus) and Eu-
ropean tree sparrow (Passer montanus) were
introduced into the U.S. in the 1850s and
1870s, respectively (Kendeigh 1973). The
house sparrow has become continent-wide,
while the tree sparrow has remained in the
general locality of St. Louis, where it was in-
troduced. As a correlary, house and European
tree sparrows were also introduced into Aus-
tralia in the 1850s and 1860s, respectively
(Frith 1977). The former species now covers
more than one-third of the continent, but the
latter has expanded to a few hundred kilome-
ters between Melbourne and Sidney. These
two examples both suggest that on the Amer-
1979
The Endangered Species: A Symposium
103
ican and Australian continents some sort of
competitive interaction or pressure which we
don't understand is being exerted that is not
operative in their native Eurasian range. We
certainly do not understand their behavioral
interactions.
If we now turn to the animal level as the
working unit, there can be, once again, an or-
ganizational breakdown for preservation or
restoration as follows (the ecosystem or com-
munity level approaches the concept of habi-
tat preservation just mentioned):
1. Species level
2. Population level
.3. Community or ecosystem level
The Species Level
Concern here may center at the local or
geographic population of the species rather
than at the species over its total range. For
example, the southern populations of the bald
eagle, which are endangered, may be of con-
siderable importance in decision making, as
opposed the northwest and Alaskan popu-
lations, which are abundant. The peregrine
falcon {Faico peregrinus tiindrius) from arctic
and subarctic regions is rare or endangered
over parts of its range, but the eastern U.S.
population (F. p. anattim) is extinct. Strate-
gies for preservation or restoration of the two
falcon populations are basically different.
The former faces reduction because of chem-
ical pollutants along the migratory route or
on its Latin American wintering grounds
(White and Cade 1977). Preservation seem-
ingly involves a land treatment practice
change in those countries. In the case of the
latter, which has already been extirpated
from its range, the strategy being employed
is the reintroduction of captive-bred stocks
into its former range. To help design the ra-
tionale and process necessary to effect a re-
covery of population numbers, as with the
falcons just mentioned, and to provide a fi-
nancial basis to help accomplish that process,
the concept of the recovery plan (Porter and
Marshall 1977, Marshall 1978) was con-
ceived.
Concern may be for the species on its
breeding grounds because those breeding
grounds may be limited or localized, rather
than for the animal at other times of the
year. For example, the northern fur seal {Cal-
lorhinus ursinus) breeds locally on the Pribi-
lof Islands, but at other times of the year it
can be widespread in the Pacific Ocean and
Bering Sea. The short-tailed shearwater {Puf-
finus tenuirostris) is localized as a breeder in
southeastern coastal Australia, but in the non-
breeding season it is very widespread and
even common.
A now common strategy at the species lev-
el is captive breeding or propagation for lat-
er release into the wild. The recent book on
endangered birds edited by Temple (1978)
covers this strategy in great detail, along with
numerous other species-oriented strategies.
Zimmerman (1975) has also chronicled sever-
al of these strategies and refers to what he
calls clinical ornithology, or the development
and implementation of techniques for specif-
ic actions.
Genetic manipulation is a strategy not yet
fully explored. The rare Edward's pheasant
{Lophura edwardsii) is being crossed with the
more common and closely related look-alike
Swinhoe's pheasant {Lophura sivinhoii). The
F-1 offspring are then Ijackcrossed with Ed-
ward's pheasant and so forth with successive
backcrosses, each time diluting the amount of
Swinhoe's genetic material. The design of
this strategy is to genetically "rebuild" the
Edward's pheasant. Wilson and Willis (1975)
carry this concept further and suggest that
certain strains of animals might be molded
genetically to fit into communities where
none now fit or where one has become ex-
tinct.
Genetic engineering (cloning, etc.) is also
now a very real possibility for the saving of
some animals by producing large numbers of
them from a single individual. Such tech-
niques, were they to become developed to
the necessary extent, could save literally hun-
dreds of dollars in the production of costly,
hard-to-breed species .such as the peregrine
falcon. On the negative side, however, gen-
etic diversity would become drastically re-
duced.
Foster parenting and cross-fostering are
other species strategies that have proven suc-
cessful with such birds as falcons and the
whooping crane (Grus americana) (cf. Dre-
wien and Bizeau 1978, Fyfe et al. 1978, and
104
Great Basin Naturalist Memoirs
No. 3
Temple 1978a).
At this time, some of the strategies that
can be devised frequently have only species-
specific application. Beyond that, the real
need is to design approaches and strategies
that will have more general and widespread
application, where several species may fit un-
der the same program in an effort to reduce
the financial burden. A list of some species,
the cause for endangerment, and strategies
projected to reduce their endangerment is
given in Table 1.
The Population Level
At this level we must first determine what
the critical or effective breeding size of the
population really is. Knowledge of that pa-
rameter may alter the approach used to help
the population. It has been suggested that the
Laysan duck {Anas laijsanensis) was reduced
to but one pair, and from there the popu-
lation has rebounded to a self-sustaining wild
population plus several score in captivity.
The Mauritius kestrel {Falco punctatus) re-
bounded from 4 individuals in the wild in
1973 to 12 in 1976 after they adopted a new
nesting habitat free from maurading monkeys
(Temple 1978b). Populations reduced to such
low levels may be subject to the in-
corporation of genetic weakness, as suggested
by Bonnell and Selander (1974) in their study
of the elephant seal {Mirounga angustriostris).
Populations that have not been reduced
Table 1. Examples of rare or endangered taxa showing known or probable major contributory causes of endan-
germent and possible or currently working strategies for preservation or trend reversal. There may be multiple
causes or strategies, but only those that are seemingly the major contributors are given. This table is meant solely to
provide examples of the array of strategies and is not intended to be complete.
Cause of endangerment
Strategy for preservation
California Condor
{Gijmnogyps califomianus)
Puerto Rican Parrot
{Amazona vittata)
Aye-Aye
{Daiibentonia madagascariensis)
Malagasy lemurs
(Lemuridae)
Night Parrot
(Geopsittlacus occidentalis)
Ground Parrot
{Pezoporus ivalliciis)
Houston Toad
{Bttfo houstonensis)
Western Swamp-turtle
(Pseitdemydiira umhrina)
Noisy Scrub-bird
(AtrichoJirnis clamosus)
Hawaiian Stilt
{liimantopiis mexiranus
knudsoni)
Socorro Isopod
{Exosphaeroma tlunnophiluni)
Masked bobwhite
{Coliniis virginiantis ridgwaiji)
Black Robin
{Petroica traversi)
Loss of habitat and food base,
disturbance, perhaps pesticides
Predation, over-exploitation, loss of
habitat
Habitat loss
Habitat loss and persecution
Habitat loss
Habitat loss
Habitat loss
Habitat loss
Habitat loss
Restricted habitat with habitat loss
Restricted habitat with habitat loss
Habitat loss or alteration
Habitat loss or alteration
Habitat purchase for preserves,
supplemental feeding, captive
breeding
Preserves, altered nesting
conditions
Habitat reserves
Habitat reserves
Habitat reserves
Habitat reserves
Habitat reserves
Protected habitat reserves, captive
breeding colonies
Protected habitat reserves
Habitat acquisition and
preservation
Maintenance of water levels,
maintenance of artificial habitat,
reestablishment into natural
habitat
Reintroduction into habitat
reserves
Reintroduction into habitat
reserves
1979
The Endangered Species: A Symposium
105
Red Wolf
{Cunis nifus)
Palila
(Psittirostra baillciii)
Hairy-nosed Wombat
{Lasiorhin us krefftii)
Bridled Nail-tailed Wallaby
{Omjchogalea fraenaUt)
P;ilinimp Killifish
(Einpctriclithys latos)
Pupfish
(Cijprinodon sp.)
Stock Island Tree Snail
{Ortlialicus reses)
Callippe Silverspot Butterfly
(Speijeria callippe)
Cranes
(Gniidae)
Whooping Crane
{Grits amehcana)
Bermuda Petrel
{Pterodroma cahow)
Giant Pied-billed Grebe
(Podihjmhiis gigas)
Utah Prairie Dog
{Cynomijs parcideiis)
Hawaiian Goose
(Branta sandvicensis)
Kirtland Warbler
(Dendroica kirthindi)
Sea turtles
(Cheloniidae and
Dennachelyidae)
Whales
(Cetacea)
Black-footed Ferret
[Mustelo nigripes)
jaguar
[Panthera onca)
Bald niis
(Gewnticus cremita)
Giant Otter
(Pteronura I)nisiliensis)
Ryuku Rabbit
(Pentalogiis furnessi)
South American River Turtle
(Podocnemis expansa)
Golden Frog
[Atelopus varius zeteki)
Peregrine Falcon
[Falco peregiinus anatum)
Arabian Oryx
(Ori/.r leuconjx)
Habitat alteration followed bv
hybridization
Habitat loss and competition
Habitat loss and competition
Habitat loss and introduced
predators
Restricted habitat, habitat loss, and
introduced competitors
Restricted habitat, habitat loss, and
introduced competitors
Habitat loss or alteration becavise
of commercial development
Habitat loss or alteration because
of commercial development
Winter habitat loss
Habitat loss, over-exploitation
Introduced and other predators,
over-exploitation
Hunting, introduced predators,
habitat loss
Over-exploitation, habitat loss
Habitat loss, over-exploitation,
introduced predators
Relict, limited habitat, habitat
alteration, nest parasites
Commercial and other over-
exploitation, habitat loss
Over-exploitation
Loss of prey base, direct
exploitation
Over-e;xploitation, persecution for
pelts
Precise causes unknown, direct
persecution?
Persecution for pelt
Predators, overliunting
Persecution for food
Collecting
Chemical pollutants
Overhunting
Secured habitat without related
can ids
Elimination of competitors from
reserves
Establishment of additional
colonies in protected areas
Protected reserves
Managed habitat reserves
Managed habitat reserves
Protected reserves
Protected reserves
Reserves, artificial feeding stations
Manipulated nesting biology, cross-
fostering young, captive rearing
Habitat preservation, nest site
modification
Habitat preservation
Relocation, habitat preservation,
legal protection
Reintroduction into habitat
preserves; captive breeding
Manage reserves, reduce parasites
Legal protection, captive rearing
reintroduction
Legal protection
Habitat and prey base
preservation, reintroduction
Legal protection from hunting
Alteration and preservation of
nesting substratum
Legal protection from hunting
Maintenance of reserves,
elimination of stray dogs
Legal protection
Legal protection
Captive breeding and
reintroduction
Captive breeding and
reintroduction, preserves
106
Great Basin Naturalist Memoirs
No. 3
below that effective size for self-mainte-
nance, although there may be few individ-
uals, can probably be maintained within
large patches of habitat or natural reserves
(Diamond 1975, Terborg 1974).
Unfortunately, natural reserves often end
up being national parks and the like. When
national parks are relegated to unused por-
tions of land or are built around scenic at-
tractions, they may fail to preserve animals in
the manner needed. Pickett and Thompson
(1978) have designed reserve concepts based
on a "minimum dynamic area" or what they
also call "patch dynamics" wherein reserves
must be large enough to maintain internal re-
colonization sources (cf. Lovejoy's comments,
this volume, on minimum critical size of eco-
systems). Such a reserve would be in effect
sort of a megazoo. These areas should have
the following properties: (1) large, (2) circu-
lar, (3) undivided or, if divided, connected by
corridors, and (4) close to one another. Figure
1 depicts a schematic drawing of such re-
serves and is based on Diamond (1976) and
Wilson and Willis (1975). Notice that nation-
al parks or monuments seldom have these cri-
teria. Hence, such reserves must be set aside
specifically for the saving of certain popu-
lations or groups of species. In dealing with
this approach in tropical rain forest in-
vertebrates, Elton (1975) has concluded that
these reserves must be very large indeed to
insure success (self-sustaining populations)
within these forests, because the organisms
are at such low densities.
The Community or Ecosystem Level
The final level at which strategies can be
aimed is the community or ecosystem level.
This is probably the level at which deteriora-
tion that eventually leads to endangerment
generally first starts. King (1978), for ex-
ample, shows that 65.3 percent of the cases
of the rare or endangered birds is caused by
ecosystem (habitat) destruction.
It is at the ecosystem level that ecological
engineering may be effective. Wilson and
Willis (1975) discuss the orphan species, or
those organisms on the brink of extinction in
their native range but capable of being fitted,
in the ecological sense, into certain alien
communities. Such "fittings" have manipu-
lative overtones characteristic of ecological
engineering. An example of such a fitting
through transplanting of a species, but not a
true orphan species {sensu stricto) in the con-
text of Wilson and Willis, comes to mind
when thinking of the Norfolk Island parrot
{Cyanoramphiis novaezelandiae cookii). This
population is unique to Norfolk Island, where
some 40 or so remain (J. M. Forshaw, pers.
comm.). A closely allied population did occur
on Lord Howe Island, but was eliminated by
persecution from early residents of the island
(both islands are in the Tasman Sea between
New Zealand and Australia). Here then, on
Lord Howe, an appropriate habitat exists in-
tact but there is no parrot to fill it. The prob-
lem of endangerment on Norfolk is the result
of the introduction of another parrot, the
crimson rosella {Platycercus elegans), an ag-
gressive, competitive species that is dis-
placing the Norfolk Island parrot. A seem-
ingly workable strategy is to transplant the
Norfolk birds to Lord Howe Island, where
they are free from another competitive par-
rot and in a habitat similar to the one from
which they were taken.
Spellerberg (1975) has worked with the
three snakes and three lizards in Britain (es-
sentially the entire British reptilian fauna),
where they are being threatened by the loss
of habitat to land development or by man-
caused fires. By examining their behavior and
ecological physiology, he concludes that the
strategy best suited to the saving of these spe-
cies is the reconstruction of islands of habitat
into which the entire reptilian communities
can be relocated.
Holt and Talbot (1978) and Wagner (1977)
have described the value of ecosystem man-
agement rather than species management.
Holt and Talbot suggest that when manage-
ment is for single stocks or species it is often
done so to the exclusion of a knowledge of:
1. Relationships within trophic levels
2. Relationships between trophic levels
3. Impact on symbiotic or commensal
relationships
4. Changes in carrying capacity due to
factors such as climate, pollution, or
other human influences
Regardless of the level at which one wishes
to approach a given strategy, much of what
1979
The Endangered Species: A Symposium
107
BETTER
WORSE
g o o o o
B
c?
o°o
o° O
Fig 1 The conceptual design of faunal reserves. These configurations are based on current theory of biogeo.
gra^h'y as IlatiTo aL of habLt, extinction, and recolonization rates. A a ^f^--J^;^^^T::^Z
peninsular effect of a long, narrow one. B: closely clumped reserves are better because of d'^^'^^^.^^f^^* ,^^^;7'^^-
connected by corridors of habitat are better than disconnected ones. D^a cont.nuous --^^ ^ ^^J^^,;*^- ^^^^^^^f
mented one because of the effect of area. (These data are modified from Diamond 1976 and W.lson and W.ll.s 1975).
108
Great Basin Naturalist Memoirs
No. 3
is done will depend on whether the goal is to
preserve or protect the animal or to restore
it, or whether the treatment is aimed at the
proximal or the ultimate cause of endan-
germent. Many of the federal or international
acts and conventions are in reality strategies
that function in providing a certain level of
protection until a plan to aid the species in
whatever way necessary can be enacted
(Schreiner and Ruhr 1974, Schreiner and Se-
necal 1978).
With the higher vertebrates most of the
endangered forms have biological character-
istics of a K-selected rather than an R-se-
lected species (Pianka 1970) and are, also, ei-
ther plagued with reduced survivorship or
reduced fecundity. These parameters were
among those considered by Adamus and
Clough (1978), Ramsay (1976), and Sparrowe
and Wight (1975) in evaluating species or set-
ting priorities for the inclusion of species into
reserves or natural areas or for program man-
agement. The priorities were given to ani-
mals based on such species characteristics
and properties as (1) suitability (mobility,
area size needs, etc.) or (2) desirability (scar-
city, endemicity, vulnerability, etc.). Ramsay
(1976) has suggested that, regardless of
whether the problem is approached through
environment (eco-unit), preservation, or the
maintenance of species diversity, a priority
system at the species level, based on econom-
ic, biological, or cultural and aesthetic values
can be developed.
Specific Strategies
Let me now cite a few specific examples
where a given strategy has been employed or
can be clearly viewed as the means to em-
ploy. The examples will apply to the species
level because most of the knowledge is there
and the problems at the population, commu-
nity, or ecosystem level are just now being
approached. These are also selected strate-
gies with simple approaches.
Example 1.: Introduced predator or com-
petition from domestic live-
stock as a major cause of en-
dangerment.
The plains wanderer (Pedionomus tor-
quatus), pig-footed bandicoot {Chaeropus
ecoudatiis), and grass owl {Tyto long-
imembris) in Australia; Aleutian Canada
goose {Branta canadensis lencopareia) in
North America; and the Galapagos tortoise
{Geochelone elephatopiis) are some examples
of these. In the case of the Aleutian goose,
the artic fox {Alopex lagopus) introduced onto
the islands nearly brought about the goose's
total demise by its predation (Springer et al.
1978). The foxes were eliminated from sever-
al islands and captive-bred geese (from the
single wild-breeding population rediscovered
in the early 1960s on Buldir Island, where
about 500 pairs breed) were then trans-
planted back onto the islands where fox had
been removed. The major obstacle now
seems to be the geese being able to make a
successful migration from their breeding
grounds to their California wintering grounds
and back to the Aleutians. The success of this
plan remains yet to be achieved. It is of inter-
est that during the decades between the
1930s and the 1970s the goose was shot on its
wintering grounds, and yet the Buldir Island
population appears to be at carrying capaci-
ty. Total protection may produce some inter-
esting changes in the nonbreeding population
structure, although all of this remains to be
documented over the next decade or so.
The Galapagos tortoises on Hood Island
were reduced to one male and 5 or 6 females
by introduced rats {Rattus sp.) that ate young
and feral goats {Capra sp.) that destroyed the
tortoises vegetative food sources (Michael
Harris, pers. comm., 1978). All the adults
were taken into captivity and about 100
young tortoises were raised. Goats have now
been removed from Hood Island (M. Harris,
pers. comm.) and the tortoises' food supply
appears to be coming back. Once the young
tortoises reach about four years of age they
are large enough to be released and rats will
not depredate them. Because the tortoises do
not breed until about 40-60 years of age,
there will be time to work on the problem of
eliminating the introduced rats. Apparently
there have been no young tortoises raised this
century on Duncan Island because of rat pre-
dation (M. Harris, pers. comm.).
In Australia there is a wingless, undes-
cribed orthropteran called P-42 (Key, 1978)
that survives in only six localized areas of
lightly grazed pastures. There they are asso-
1979
The Endangered Species: A Symposium
109
ciated with one of the native composite
plants of the genus Helichnjsum. This com-
posite is ehminated by heavy Hvestock graz-
ing. The apparent strategy is a simple one of
controlling or eliminating grazing from these
localities. If some of these localities are on
Crown Lands, the task of controlling grazing
will be made easier.
Most populations on islands (either habitat
or geographic islands) can be protected by
simply controlling the introduction of pre-
dators or by eliminating them from islands
where they have previously been introduced.
Many of the examples in Table 1 fall into this
category.
Example 2.: Major habitat alterations or
losses as a principal cause of
endangerment.
A prime example in such a case can be
seen in the bony-tail chub {Gila elegans), ra-
zor-back sucker {Xijmuchen texanus), and
Colorado squawfish {Ptychocheihis hicius) in
the Colorado River. The construction of
dams or direct use of the water by man has
caused at least two major classes of change,
namely, (1) alterations of natural water cycles
(dam, dewatering, stream flow changes, etc.)
and (2) water quality changes (silt loads, tem-
perature, pollution, etc.) (Seethaler 1978,
Minckley 1973). The introduction of exotic
fishes may also have helped to reduce these
species to critical levels. Breeding cycles,
correlated with such parameters as silt loads,
stream bottom morphology, and temper-
ature, have been reduced or eliminated and,
although attempts at spawning by the chub
and squawfish are observed, juveniles are in-
frequent or lacking. A nice correlary exists
with the Macquarie perch {Maequoria aiis-
tralasica), trout cod (Maccullochella mit-
chelli), and Murray cod {Maccullochella mac-
quariensis) of the Murray-Darling river
system in Australia. These fish need high silt
loads, flooding, and fluctuating river condi-
tions, as are common with spring rains, to in-
itiate spawning (Lake 1971). Dams and reser-
voirs have flattened out fluctuations and
lowered silt loads. On one portion of the riv-
er there has been no flooding since 1939 be-
cause of dams (Roughley 1951). The fisheries
industry and introduced European carp
{Cyprinus cai-pio) have also had their impact
on reducing these species. One solution to
this problem in both North America and Aus-
tralia is to introduce these species into
streams with the neces.sary parameters where
dams do not occur. Currently, of about 50
species of Australian fresh water fishes (most
are endemic), about one-third are endangered
or threatened, many because of man-caused
changes in water conditions (Lake 1971).
The above examples tend to be very
straightforward causes of endangerment, and
in some cases the solution to alleviating the
problems is rather simple. Most frequently,
however, there are not single but multiple
causes of endangerment. These have no easy
or readily obvious workable strategies for sol-
ving the problem. The Higgins eye mussel
{Lampsilis higginsi) of the Mississippi River
system is an example wherein at least the fol-
lowing six factors have a measurable adverse
impact:
1. Excessive commercial exploitation
2. Water quality degradation (industrial
wastes, pesticide run off, etc.)
3. Increased siltation and turbidity
4. Dredging
5. The effective impact of exotic clams
which dislodge the Higgins eye from at-
tachment locations
6. Possible reduction of host fish species
for larvae
As a final statement, perhaps the most im-
portant and critical strategy at this stage is to
engage young, innovative minds in determin-
ing quickly and efficiently what the problems
are, wed the multiple interests and groups
into a common cause, decide what the prior-
ities should be, and solve the specific prob-
lem at a minimum effective financial cost.
Something as simple as convincing the Aus-
tralian farmer that dead snags of the river red
gum {Eucalyptus camaldulensis) do not take
up significant space and that he should leave
them in his field may be all that is needed.
These trees contain many cavities and pro-
vide nesting places for a myriad of species.
That one simple judgement, not to burn
down the dead tree, may have far-reaching
impact. Ramsay (1967) has rightly pointed
out that our judgments must be preceded by
as much forethought and rationality as pos-
sible because developing priorities for preser-
110
Great Basin Naturalist Memoirs
No. 3
vation decisions favors an analysis of species
from the viewpoint of human values. These
values may indeed not be the critical ones for
the integrity of a healthy ecosystem. None-
theless, some value judgements (Myers 1976
and Holliday 1978) as to the worth of species
and man's stewardship must be the first step
in the long and never-ending process of
maintaining a diverse ecosystem.
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1979
The Endangered Species: A Symposium
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RARE SPECIES AS EXAMPLES OF PLANT EVOLUTION
G. Ledyaid Stebbins'
.\bstract.- Rare species, including endangered ones, can be very valuable sources of information about evolution-
arv processes. They may be rare and valuable because: (1) they are evolutionary youngsters and could represent an
entirelv new evolutionary strategy of great scientific and practical value; (2) they are evolutionary relicts that have
stored enormous amounts of genetic information of great worth; (3) they may represent endemic varieties that har-
bor a great deal of the genetic variability in the gene pool that would be of enormous value to a plant geneticist; the
rarity of the plant is not necessarily correlated with the size of its gene pool; (4) they may represent unique ecologi-
cal adaptations of great value to future generations. Studies of gene pools and the genetics of adaptation constitute a
new and developing field of the future.
This morning I'm talking about a some-
what different topic than yesterday evening
and one more in hne with my usual interest
because, as many of you know, I am primari-
ly an evolutionist. The theme that I would
like to develop is that rare species, including
the endangered ones, can be very valuable
.sources of information about evolutionary
processes. To illustrate that theme, I am go-
ing to give you examples of four alternative,
but not mutually exclusive, theories that have
been suggested for the reason that species are
rare. One of the oldest, which was cham-
pioned in the early part of the 20th century
by J. C. Willis, was that the rare endemic
species were youngsters. They appeared on
the earth recently and haven't had time to
spread. He wrote a whole book on the sub-
ject, which he called Age and Area. I first
heard of that book from my systematics pro-
fessor, M. L. Fernald, who was very much in-
terested in rare species. He regarded them as
senescent relics, and strongly opposed Wil-
lis's theory. However, Willis was partly cor-
rect. We have perhaps more direct evidence
of this than for any other hypotheses. This is
expected, because the origin of some new
species would be recent enough that we
would know when it appeared. A now classic
example is that presented by the late Marion
Ownbey, of Washington State University,
with respect to the goatsbeard genus, Trag-
opogon.
Projected on the screen is a chart which
Ownbey put in the American Journal of Bot-
any in 1950. The three species which are list-
ed all have the chromosome number 2n= 12.
They are well-known European plants in-
troduced as weeds into North America.
There are no species of Tragopogon in the
Old World that have chromosome numbers
higher than 2n=12 and n = 6. However, in
the backyards and railroad yards around Pull-
man, Washington, Ownbey found two differ-
ent entities having 24 chromosomes, twice
the number of all the others in the genus. By
a series of ingenious hybridizations and analy-
ses of chromosomes, he established without
doubt that one of them, described as a new
species, T. miscellus, was derived from hy-
bridizations between T. ditbius and T. pra-
tensis. Its chromosome number has been
doubled either before or after the initial
hybridization, giving us a stable intermediate,
a principle long known to geneticists. A sec-
ond species, T. minis, has originated in the
same way from a cross between T. porrifoUus
and T. duhius. Because T. dubiiis did not ex-
ist in western North America before about
the 1890s, the age of these two species when
he first found them was not more than a half
century. They now are spreading. They are
known in one or two places in Montana.
There is another locality in Arizona. They
are certainly not endangered. A hundred
years from now, they may have become com-
artmenl of Genetics, University of California. Davis, California (
113
114
Great Basin Naturalist Memoirs
No. 3
mon weeds.
There are other very young examples, not
quite so young as these. Indirect evidence
suggests that some species have arisen since
the retreat of the glacial ice from northern
North America and the drying up of the cli-
mate in places like California where the
Pleistocene was a rainy or pluvial period.
One example is that of three species in the
genus Laijia, or "tidy tips," an annual plant
of the composite family, or Asteraceae. The
three species involved are L. jonesii, L. mun-
zii, and L. leucopappa. They are very closely
related, on the l3order line of becoming spe-
cies. They can be hybridized easily and the
hybrid is partly fertile. They are more dis-
tantly related to L. fremontii and L. platij-
glossa.
Their distributions are as follows: Laijia jo-
nesii occurs near the coast of south central
California, not far from San Luis Obispo;
Laijia miinzii is found in the small valleys of
the inner south coast ranges; and L. leuco-
pappa, in the central valley of California, on
one or two hillsides not far from Bakersfield.
They are closely related entities occupying
neighboring areas in the same general region.
The climate of this region was drastically
changed in the Pleistocene. It seems likely
that the splitting of these three populations
from each other is post-Pleistocene, about six
to eight thousand years ago.
Layia miinzii is on the border line of being
rare in a dry season, but is certainly not en-
dangered. Layia jonesii, being on the coast,
might be endangered. Layia leucopappa, in
the highly cultivated central valley, definite-
ly is endangered. A nature conservancy
group, located in Bakersfield, is very much
interested in it. It occurs on a large private
range area, the manager of which says they
are going to preserve it for us. This is a case
of a young species which is on the rare and
endangered list.
Another example of a similar nature is a
species of larkspur. Delphinium hesperinm is
a common widespread species of the oak
woodlands in the inner coast ranges of Cali-
fornia. Delphinium recurvatum is foimd in
the central valley and bottom lands and Del-
phinium gypsophilum, as the name implies,
lives in gypsumlike soil on the hills bordering
the valley. It occupies a habitat between the
other two species. Delphinium hesperinm and
D. recurvatum have been crossed. The hybrid
is partly fertile and looks like D. gypsophi-
lum. The interesting thing is that when the
F-1 hybrid is crossed with native D. gyp-
sophilum it is found to be more compatible
than the cross with either of its own parents.
This is pretty good evidence that the native
D. gypsophilum is derived from hybridization
between the two other species. This again is
a recent species. It is too common to be on
the rare and endangered list, but it is cer-
tainly recent and uncommon relative to its
ancestors.
An ancient species is the California Big
Tree, Sequoiadendron giganteum. It is con-
fined to certain groves in the Sierra Nevada
in California, extending from Sequoia Na-
tional Forest in the south in Tulare County
into ever-smaller groves, to a very tiny one in
the northernmost area in Placer County to
the west of Lake Tahoe. It does not have a
much-restricted gene pool. Horticulturists
have found they can extract various modifi-
cations of it by simple inbreeding. Daniel Ax-
elrod, with his brilliant studies of the pa-
leobotany of western Nevada, has shown that
California Big Tree was very abundant in
Pliocene forests, six, seven, or eight million
years ago in western Nevada when the Sierra
Nevada didn't exist and the climate of Ne-
vada, due to moist winds from the ocean, was
still relatively mesic. Because of the Pleisto-
cene changes in climate, it has become re-
stricted in two ways: (1) the elimination of
the stands to the east of the Sierra Nevada by
aridity and (2) the reduction of the stands in
California by the increasing length of the
rainless dry summer. I say this because of re-
search by the late Woodbridge Metcalf, ex-
tension forester at Berkeley. He showed that
the big tree would seed itself only in a year
when the dry season from May till October is
shorter than usual. If it gets a few of these
short dry seasons between seed germination
and the time when the trees are 8-10 years
old, then it competes with sugar pine, white
fir, and other forest species. If young trees
are exposed to a succession of normal dry
summers, they cannot compete with the
seedlings of the other species. This, then, is
an example of an ancient species.
There are some others. One of the inter-
1979
The Endangered Species: A Symposium
115
esting things that Professor Jack Major and I
worked out some years ago is that the con-
centration of rare rehctual species, which
were more common in the past, is bipolar in
Cahfornia. It represents two elements, first a
northwestern element which is related to the
Pacific Northwest and Asia. The species in-
volved are mainly trees or shrubs related to
holarctic members of the same genera. Exam-
ples are the Weeping Spruce {Picea brewe-
riana), which is narrowly restricted to this
area. The Sadler Oak {Qiiercus sadleriana) is
related to the Chestnut Oaks of the eastern
United States, and is restricted to the Sis-
kivoii Mountain area. The Port Orford Cy-
press (Cliamaecijparis lawsoniana) is in this
area. It contains several endemic species of
mesic genera like Vancouveria and some gen-
era of the saxifrage family.
In the southern area occur rare, endemic
species like Hesperocallis of the Lily family,
the jojoba shrub, Simmondsia, not particu-
larly rare but certainly relictual.
Bv contrast, central California contains en-
demic species, which seem to be new like
those of ArctostapJnjlos or Manzanita, al-
ready discussed, and other annuals. Because
central California has received the greatest
disturbance in relatively recent times, one
can get some idea of whether a species is re-
lictual or not, both by its taxonomic affinities
and by the geographic areas in which it
grows.
An example of an herbaceous species from
the northern relictual area is Darlingtonia
caUfornica. It does not have official preserva-
tion that I know of in California. It should,
because it is an attractive species, an insect-
digesting pitcher plant. Every high school bi-
ology teacher wants one in the classroom,
and people also like them in their homes.
There is danger to them from vandalism. I
sometimes blush when I go up the Oregon
Coast Highway and see that Oregon has a
preserved area of Darlingtonia labeled as
such, while California doesn't.
The next group of rare and endangered
species are of an entirely different nature.
Here, I want to state publicly that one of my
articles written in 1942, and often quoted,
apparently is not correct. In 1942, I made the
speculation that rare and restricted species
would usually be so because of restricted gen-
etic diversity. At that time I was influenced
by the work of Sewall Wright on inbreeding
and its ramifications, which was very popular
at that time. Now we have data on restricted
species like Clarkia franciscana, which grows
in only a single hillside in the city of San
Francisco and which, as my colleague Leslie
Gottlieb has shown, has as much biochemical
variability in it in terms of enzyme alleles as
Clarkia ruhicunda, which is much more
widespread throughout the San Francisco
Peninsula. I now reduce the category of spe-
cies that are rare because of restricted gene
pool to a rather small one consisting of spe-
cies that have not only become inbred be-
cause of small population size, but, in addi-
tion, because of the shift from cross-
fertilization to self-fertilization or pre-
dominant fertilization. Again, Leslie Gottlieb
has shown with his work on enzyme varia-
bility that in eastern Oregon there is a wide-
spread species of the composite genus Steph-
anomeria {Stephanomeria coronaria), which
has an enormous amount of genetic varia-
bility, but right next to it there is a very re-
stricted species along Malheur Lake in east-
em Oregon, Stephanomeria malheurensis,
which has very little variability. One differ-
ence between the species is that S. coronaria
is largely outcrossed and S. malheurensis is
almost completely inbred. The inbreeding as
much as the restriction in size of the popu-
lation and habitat was responsible for the re-
duction in size of the gene pool. The same
situation exists in some species of animals like
the burrowing rodents of the Middle East,
studied by several Israeli zoologists, and cer-
tain fishes in Mexican caves which have very
restricted gene pools. One that is rather
widespread, the southern alligator, apparent-
ly has extremely low variability in its gene
pool. The whole hypothesis that there is a
strong correlation between rarities and en-
dangeredness and the size of the gene pool
must be greatly modified, if not entirely re-
jected. In my own thinking, I am substituting
the concept of ecological traps. Plant species
may be hemmed in by environments that are
so different from the ones in which they
grow that they do not have the genetic po-
tential to colonize those habitats. Sometimes
the "traps" are quite clearly defined so that I
call them ecological islands. Usually the soil
116
Great Basin Naturalist Memoirs
No. 3
is so different on the islands, relative to that
of the surrounding islands, that the plants are
as if they were on an oceanic island sur-
rounded by a sea of unfavorable soil condi-
tions. Such an island is Pine Hill, 25 miles
east of Sacramento. It consists of basic intru-
sives, a very distinctive type of rock, sur-
rounded by the various metamorphic rocks
commonly found in the Sierra Nevada foot-
hills. Interestingly enough, this particular is-
land is believed to have once been an oceanic
island. Dr. Eldredge Moore of our geology
department at Davis, in association with the
new theory of Plate Tectonics, points out
that basic intrusives are associated with the
roots of volcanos. The situation could be ex-
plained if the rocks now exposed at Pine Hill
were the roots of an ancient submarine vol-
cano that arose in the Pacific Ocean to the
west of what then was the seashore, but is
now the eastern edge of the central valley.
Because of cnistal movements, this ancient is-
land jammed itself against the older rocks.
A striking endemic on Pine Hill is Fe-
montodendron californicum ssp. decumhens,
a prostrate shrub that bears highly distinctive
copper-colored flowers.
Another ecological island is in the area by
Monterey and Pacific Grove. There are two
species of rare endemic cypresses of the area.
Neither of them is in danger now, because
both are preserved. One is the well-known
Monterey cypress {Ciipres.sus macrocwpa),
found only on the granite ledges near the
shore. The other is the Gowen cypress, found
only in the hard pan of the raised beach, in
the interior of the peninsula.
Monterey pine is confined to the Monterey
ecological island plus two others in Califor-
nia, one 50 miles north, the other 120 miles
south. There is evidence that it is not restrict-
ed in its colonizing ability, if it has the right
conditions, so that it could get out of its eco-
logical trap. This is evident from what the
Monterey pine has done in the southern hem-
isphere, in Chile, New Zealand, and Austra-
lia. In all three regions, extensive forests of
this species have been planted that in many
places look quite natural. Some trees are far
taller than those in California, reaching
heights of 100 to 150 feet.
Most interesting of all, in the vicinity of
Canberra, Australia, I have seen Monterey
pine seedlings invading a forest of native Eu-
calyptus.
A well-known ecological island exists in
the Sierra foothills of California, near the
town of lone. It is based on a hard pan soil
which is of Eocene age, about 40 million
years old. It was a sea-beach terrace facing
the Pacific Ocean, which at that time cov-
ered all of the present central valley. It is
dominated in many areas by an olive-colored
shrub, Arctostoplujios mijiiifolia, the lone
manzanita. There is a margin of the common
gray manzanita, Arctostoplujios viscida, and
interior live oak, but most of the area sur-
rounding it is grassland consisting of in-
troduced annual grasses and scattered blue
oaks and digger pines that are the normal
dominant vegetation of the area. The plants
that are rare and endangered are held in
check by the very special ecological condi-
tions that prevail here. There is really an is-
land within an island because a buckwheat,
Eriogonum apricum, which is confined to the
lone manzanita island, grows only in the few
most barren parts of it.
The gene pool of E. apricum is most inter-
esting. There are three patches of it. The
maximum distance from one to another is
about 10 miles and there is one about half-
way in between. Rod Myatt of UC Davis did
a master's study on morphological variability
and found that the different patches can be
distinguished morphologically. They are
races. There is not only a lot of variation
within each of these patches, but distinctive
differences between patches. In other words,
it appears as if the lone manzanita within its
area of 10 miles has as much morphological
variability as does another buckwheat,
Eriogonum nudum, within an area of equal
size, 10 miles in diameter in the Sierra foot-
hills. The difference is that Eriogonum nu-
dum, one of the most common buckwheats in
California, has a multitude of races that are
adapted to all sorts of climatic conditions
over this extensive area. Another fact is that
E. nudum is a much bigger species and its
seeds are much bigger. Its seedlings are prob-
ably much more competitive so that it could
colonize new areas more easily than could
Eriogonum apricum. Perhaps E. apricum is a
relic of the days when annuals, even native
annuals, were much fewer than they are now.
1979
The Endangered Species: A Symposium
117
It may have colonized the barren places that
no other perennials could live in, when there
wasn't so much annual competition. That's
pure speculation. This again emphasizes the
ecological entrapment which I believe is the
basis for imderstanding the distribution of
most of our rare and endangered species, in-
dependent of age and independent of quan-
titative sizes of gene pools.
The final example is similar. It is Convict
Canyon in the eastern Sierra Nevadas. It is
distinctive, because while most of the Sierra
is either granitic or ancient acidic crystalline
volcanics of Mesozoic age or earlier. Convict
Canyon has a vein of limestone running
through it. It is the only sizable part of the
Sierra Nevada that has limestone in associ-
ation with alpine or subalpine conditions.
Mount Baldwin, 12,000 feet high, supports
a rare rock cress, Draba nivalis, and is the
only locality in the Sierra Nevada. The near-
est to it, as far as I know, are the Ruby
Mountains in eastern Nevada.
One of the most remarkable plants, how-
ever, in this area, is a willow which is related
to a Rocky Mountain species. Salix brachyo-
carpa. Another species is a relative of the
sedges, perhaps ancestral to the genus carex
Kobresia mijosuroides and an extremely local-
ized dwarf bullrush, Scirpus rollandii, some-
times put in the species Scirpus pumilus,
which has been the subject of interest for arc-
tic alpine botanists for many years. The two
discoverers of these species were Jack Major
and Sam Bamberg.
Now, why do these rare plants grow here?
It isn't because they are lime-loving calco-
philes. The Kobresia grows on Mount Evans
in Colorado which is acidic granite. Certainly
Scirpus rollandii is to a certain extent a cal-
ceophile, but it doesn't seem as though the
limestone is the basic reason. The other fac-
tor is this— limestone is porous. Because it
holds water, on a steep slope like the one
which supports the rare plants, water oozes
out from the ground throughout the summer.
Remember that the Sierra Nevada, in con-
trast to the Rocky Mountains or the Wasatch
Range, has very few summer storms. They ex-
ist, but they're small and most of them hardly
wet the ground. Much of the sierran area in
the well-drained slopes becomes very dry in
the summer, and the mesic plants have to
grow where they get heavy snowfall during
the winter. The rare plants, however, grow
on a bench area that gets relatively less snow
during the winter which is constantly coming
out of the limestone formation. This, I be-
lieve is responsible for the unusual nature of
the environment.
My final remark has become obvious from
what I've had to say. Ecological genetics is a
relatively new field. The combination be-
tween studies of gene pools and the genetics
of adaptation, I predict, is a field which is
just beginning to emerge and will be explo-
sive in the next half century. Young scientists
who are interested in native environments
and wish to study them in depth from an ana-
lytical point of view will have an exciting ca-
reer of discovery ahead of them.
THE MEANING OF "RARE" AND "ENDANGERED"
IN THE EVOLUTION OF WESTERN SHRUBS
Howard C. Stiitz'
.\bstract.— In the evolutionary process, species continually come and go. Consequently, all species on earth
today were, at one time, "rare and endangered" while in their infancy, and most will become "rare and endangered"
once more as they are replaced. Therefore, decisions relative to protecting rare and endangered species are largely
meaningless if based on numbers alone. They must include information about their biology and evolutionary history.
Lists of endangered forms currently being prepared apparently include ouly those which are (1) scarce (rare and of
restricted distribution), (2) named, and (.3) sponsored. Their biological, economic, and academic values may be more
important, but apparently are not often considered. As abundantly illustrated in western shrubs, genetically rich
genotypes are sometimes maintained by only a few individuals, whereas uniform, and therefore rare, genotypes may
in some circumstances, be represented by many individuals in uniform environments. Wise management decisions
cannot, therefore, come from numbers alone.
Interpretations of the origin of species in-
dicate that all species now on earth were at
one time rare and endangered. Whether they
arose slowly by accumnlating mutations that
permitted divergence from parental forms,
explosively as polyploid derivatives, or as re-
combinants from interspecific hybrids, they
all had humble, precarious beginnings. Fur-
thermore, they represent but a tiny fraction
of all that might-have-been. Many are un-
doubtedly inferior to former contempo-
raneous taxa which, although superior gen-
etically, were lost by fortuitous accidents
during their infancy.
As species come and go in response to the
challenge of an ever-changing world, some
are rare simply because they are new, others
are rare because they are being replaced by
more adaptive competitors. All species are
endangered in the sense that they are success-
ful only as long as the environment in which
they are superior endures, or until other
modified, improved competitors replace
them.
Intelligent intervention in this efficient, .sif-
ting, ever-improving drama in the guise of
protecting threatened species, requires there-
fore understanding the evolutionary dynam-
ics which define them. Because artificial pro-
tection of any species may concomittantly
impose intensified selection against all other
associated species, utmost care and caution is
essential in management decisions designed
to deliberately favor specified taxa. Some
species, represented by many individuals but
which are genetically uniform, in certain cir-
cumstances may be far more in danger of ex-
tinction than "rare" species which are gen-
etically diversified.
Protective measures aimed at preserving
one particular taxon may be detrimental to
the entire ecosystem. However, rare forms
which are of high intrinsic value because of
their potential for improving an ecosystem,
or for providing a fountain of genetic varia-
bles from which other new improvements
can arise, or for providing economic or aes-
thetic benefits for mankind may deserve
deference and intense protection.
Decisions regarding management of eco-
systems designed to preserve "rare" and "en-
dangered" species are therefore always pre-
carious and are essentially indefensible unless
founded on intimate knowledge of the gen-
etics and genealogy of affected species.
As rosters of rare and endangered species
begin to emerge, it is important that defin-
able criteria be used for deciding whether or
not a species is to make the roster. Apparent-
ly, to date, only three ingredients are re-
quired:
'Department of 1
and Range Science, Brigham Young University, Provo, Utah i
119
120
Great Basin Naturalist Memoirs
No. 3
(a) Scarcity (rare and of restricted distribu-
tion)
(b) A name
(c) A sponsor
Apparently it has had nothing at all to do
with value. Also, (b) is not independent of (a)
nor is (c) independent of (b). If scarce, a spe-
cies may not have a name; if unnamed, it will
almost never have a sponsor.
Actually, however, because favoring one
species may concomitantly disfavor another,
decisions cannot really evade value judge-
ments. In my opinion, they should not. I
would recommend that they deliberately in-
clude at least the following:
(a) Aesthetic values, including beauty,
uniqueness, antiquity, etc.
(b) Biological values, particularly in rela-
tionship to genetic potential and con-
tributions to the ecosystem.
(c) Economic values which would include
their contribution to wildlife, to range
use, to industry, to recreation, etc.
(d) Academic values including contribu-
tions to the interpretations of evolu-
tionary history, geological events,
climatological changes, and ecological
succession.
Illvistrations of rare forms which are being
replaced and may, therefore, not invite hu-
man intervention for their protection, as well
as forms which are "brand-new," exciting,
promising, arrivals and may, therefore, prof-
itably be enhanced, are abundant in western
North America. Recent major geological and
climatological changes have provided a mul-
titude of new habitats in which newly formed
species have been and still are being favored.
Concomitantly, similar other populations
have become reduced or extinguished as their
required niche disappeared. Examples may
be found in nearly all groups of plants and
animals; the following are illustrative:
1. The Rose Family
In the rose family, Cercocarpus, Purshia,
and Coivania have all shown explosive re-
sponse to recent habitat changes. However,
management decisions concerning the ac-
companying rare and endangered forms of
these genera will of necessity vary, simply
because each has a distinctive evolutionary
meaning.
In Cercocaijms, three principal species are
known in the Great Basin: C. montaniis, C.
ledifolius, and C. intricatus. Natural hybrids
are common between C. ledifolius and each
of the other species, but are rare between C
montanus and C. intricatus, even when they
are sympatric (Plummer et al. 1957, Pyrah
1964). Cercocai-pus intricatus is the most xer-
ic of the three, often growing on steep, ex-
posed limestone cliffs, but it is found only in
Utah and the immediate borders of neighbor-
ing states. In areas where C. intricatus and C
ledifolius grow together, there is often such a
continuum of intermediates that individual
plants are difficult to define. For these rea-
sons, it appears that C. intricatus has been re-
cently derived from C. ledifolius, having ac-
quired adaptive attributes by rapid genetic
assimilation of drought-resistant phenotypes
which were, and still are, latent in C. ledi-
folius. Similar evidence, although less
straightforward, suggests that C. montanus
may also have been derived from C. ledi-
folius by selection of types that were more
competitive in the more densely vegetated
mountain brush zone. The requirement for
broader leaves, an apparent prerequisite for
competition with Quercus gambellii, Ayne-
lanchier alnifolia, and Prunus melanocarpa,
was apparently possible only if these broad
leaves also became deciduous to escape the
long winter drought of frozen soil.
If these interpretations of the recent and
continuing evolution of Cercocarpus are
valid, what might our decisions be, relative
to the component rare and endangered
forms? Cercocarpus intricatus, although re-
cently derived and somewhat rare, is cur-
rently not endangered and probably needs
little, if any, artificial protection. Its habitat
is not often used by man nor by domesticated
animals. Very little of the current impact of
human civilization appears to be in any way
threatening this species.
Hybrids and hybrid derivatives, however,
are a different story. Not only are they very
rare but they are also very important reser-
voirs of potential diversity and, in many
cases, severely threatened. They are of value
as demonstrations of biological evolution, as
fountains of genetic combinations from
which both C. montanus and C. intricatus
might be enriched and from which even oth-
1979
The Endangered Species: A Symposium
121
er species might arise, and as beautiful, rare
specimens, with simple intrinsic aesthetic
value. Although these populations may not
represent species, they are important and in
many instances deserve deliberate protection.
However, because they are unnamed and,
perhaps, even unnamable, because the indi-
vidual plants are the unique rare entities,
they may never make the roster.
Furshia and Cowania, two other genera of
the rose family, are also rapidly evolving and
have recently produced several new adaptive
products (Stutz and Thomas 1964). While
some of the new forms may have come from
new mutations, almost all appear to have
come as adaptive segregation products from
intergeneric hybrids between them.
Purshia tridentata is distributed from
southern Utah northward into Canada. Cow-
ania stanshiinjana grows from northern Utah
southward into Old Mexico. Consequently,
almost the entire state of Utah is an overlap
zone in the distribution of these species. In
many places in Utah, where these two species
come together, hybrids and hybrid deriva-
tives are common. So commonplace is such
hybridization there appear to be no popu-
lations of Purshia tridentata in all of Utah
which do not contain introgressants from
Cowania (Stutz and Thomas 1964). Many,
perhaps most, are one of a kind. They appar-
ently continually come and go with few if
any ever being superior to their progenitors.
Here then is an example of species in
which rare and endangered forms are ramp-
ant. But, as interesting as they are or as po-
tentially valuable as they may be, attempts to
protect them all would be absurd. Selected
forms, or even selected individuals, may be
locally desirable, but it would be impossible
to preserve every noble segregant. The Cow-
ania X Purshia Fj hybrid and the segregat-
ing hybrid swarm northeast of Provo, de-
scribed in detail by Stutz and Thomas (1964),
might have been profitably spared but, be-
cause they have both already been com-
pletely obliterated by a recent housing devel-
opment, that is now impossible. Protection of
similar known F^ hybrids and hybrid segre-
gants is probably unwarranted. However,
specific products that show unique adaptive
promise may be profitably protected.
Near Clarkston, Cache County, Utah, a
distinctive population derived from Coivania
X Purshia parentage is on the verge of being
exterminated by overgrazing. Although con-
siderable segregation is still evident, many of
the plants appear to be stabilizing around a
unique combination of characteristics which
are apparently adaptive in this area. The
fniits, leaves, and habit are all intermediate
between their putative parents. Because it is
unlikely that this small population will sur-
vive long under present grazing pressures,
adjustments in management of this area
would seem highly desirable. However, it is
unnamed and unnamable and will probably
not make the roster even with me as its spon-
sor.
Other adaptive products from this parent-
age include a species of recent origin, Purshia
glandulosa Curran, and a series of popu-
lations of Purshia tridentata throughout
Idaho, Oregon, and Montana that have been
enriched with Cowania genes by in-
trogressive hybridization. Because the in-
trogressed populations are being differen-
tially favored by current grazing practices,
they apparently require no deliberate pro-
tection. Sheep apparently prefer Purshia tri-
dentata plants that contain no Cowania
genes, so introgressed populations are becom-
ing increasingly abundant. These "rare"
forms are therefore not at all endangered and
may eventually prove to be very abundant
and perhaps even detrimental as range for-
age. In this case, the rare does not at all
equate with endangered. Already it is becom-
ing difficult to find "pure" nonintrogressed
populations of P. tridentata. In time, they
may indeed become the rare and endangered
representatives.
Each population of P. glandidosa is also
somewhat genetically unique. The particular
combination of Cowania and Purshia genes
that identifies the adaptive features of P.
glandulosa is common to all populations, but
other characteristics, under less severe selec-
tion, apparently segregate somewhat ran-
domly. Consequently, plants in every popu-
lation are similar with respect to the unique
features which characterize them as P.
glandulosa, but they differ considerably in
other segregating attributes. Already P.
glandulosa as a taxon is sufficiently well es-
tablished that it is far from being rare or en-
122
Great Basin Naturalist Memoirs
No. 3
dangered. Although individual variants may
be "rare," they appear to be of little con-
sequence in the evolutionary drama that is
producing P. glandidosa as a newly derived
species. Consequently, there appears to be
little wisdom in deliberately preserving them
even though they meet the rare and endan-
gered criteria.
2. The Oaks
Much of the variation in Quercus gambeJii
in northern Utah appears to be the result of
introgression from Q. turbinella. Although in
Utah these species are currently sympatric in
only a limited area in the southern part of
the state, hybrids are common along the
Wasatch Front 200 miles to the north (Cot-
tam et al. 1959). According to those authors,
the hybrids were apparently left behind dur-
ing the altithermal postglacial period when,
because of milder climates, Q. turbinella was
able to grow that far north. Although most of
the intermediate forms are much alike and
mav be mostly Fj hybrids, some segregation
is apparent. Both "Fj hybrids" and segregants
are severely restricted to a narrow temper-
ature-inversion belt at about 5,400 feet eleva-
tion where temperatures are normally higher
than either above or below (Cottam 1959).
These rare hybrid specimens are of high
aesthetic value and apparently also of high
biological significance. If, as appears likely,
much of the expressed variation in Q. gam-
belii is due to introgression from Q. tiirbmella
via these hybrids, they have already made
great biological contributions and promise to
continue to do so as long as we permit them
to remain.
However, many of these valuable speci-
mens have already been destroyed and most
of those that remain are threatened with ex-
tinction. One very unusual hybrid derivative
near the mouth of Immigration Canyon, east
of Salt Lake City, Utah, has recently been re-
placed by a house. This particular plant
differed from both parents and all other seg-
regants in having oval leaves with serrate
margins. The leaves resembled superficially
those of chokecherry {Primus melunocm-pa).
It was a magnificent specimen with a speed
of about 40 feet. It should have been pre-
served. If anything can be done to save the
others, they will have longlasting significance
biologically, aesthetically, and economically.
But these unusual plants {Q. X pauciloba)
may never make the roster.
3. The Saltbushes
Many of the new habitats which have re-
cently become available in North America
are still completely vmoccupied. Species have
not yet evolved that can accommodate the
niunerous steep mud hills, salt flats, and alka-
li playas that characterize much of the west-
ern deserts. The plants at the borders of these
sterile islands, and therefore closest to in-
vading them, are almost all members of the
family Chenopodiacea: Salicornia, Allen-
rolfia, Suedo, Sorcobatus, Salsolo, Hologeton,
Graijia, and Atriplex. Most of these genera
are represented by only a few species and are
therefore probably there because of charac-
teristics acquired elsewhere that made them
preadapted to these harsh sites. The principal
exception is Atriplex. This genus is represent-
ed by numerous species and varieties, many
of which appear to be of very recent origin.
In some cases new successful forms appear to
be no more than a few years old.
Every known evolutionary force appears
to be operative in Atriplex at an accelerated
rate (Stutz 1978). Species appear to be aris-
ing from new mutations, from introgressive
hybridization, as new hybrid segregants from
interspecific hybrids, as autopolyploids, and
as allopolyploids. Rare and endangered forms
are therefore abundant. Some are of obvious
high value; many others are undoubtedly im-
portant.
One of the most successful species of Atri-
plex to invade western North America is A.
conescens (fourwing saltbush). It has the
widest distribution of all native perennial
species, growing from central Mexico to Can-
ada and from the Dakotas to the Pacific
Coast. With such a wide distribution, it is
probably not surprising that it is a highly var-
iable species. Some of the variation is due to
phenotypic plasticitv, but most of it appears
to be genetic.
Four different chrosome levels in Atriplex
canescens are known: diploids, tetraploids,
hexaploids, and twelve-ploids. Rare and en-
dangered forms are foimd in each.
A. The diploids (2n = 18)
Individual diploid plants have been found
1979
The Endangered Species: A Symposium
123
sporadically in several polyploid populations
and hence are probably derived by poly-
haploidy. They are certainly rare, and cer-
tainly endangered. But should they be pro-
tected? Probably not. None appears to have
any capacity for increasing (partly because,
being rare in a polyploid population, they
can leave only sterile offspring). With more
knowledge, some of them might be recog-
nized as potentially valuable entities and
might therefore warrant careful propagation
and ultimately increase for some specific use.
For the most part, however, we might expect
these to be continually produced and contin-
ually discarded as novel but nonadaptive var-
iants. Being rare and endangered in this case
is probably insufficient license to receive any
special protection.
Three distinctive diploid varieties are
known, however, which are highly successful
in specific habitats and are therefore very
valuable. At least one of them is sufficiently
rare to be considered endangered. All are
probably relics derived from ancestral dipl-
oids rather than polyhaploids derived from
polyploids.
The most abundant of these diploids, and
probably the most ancient, is a form which is
common in southern Arizona and also report-
ed from southwestern New Mexico by Max
Dunford (oral comm.). It appears to be the
most drought resistant of all forms of A. ca-
nescens, growing sympatrically with creosote
bush {Larrea tridentata) and mesquite (Proso-
pis glandulosa).
The other two diploids are narrowly en-
demic. One (A. ganettii) grows only in loose
sandstone-talus along the Colorado River and
at the mouths of its tributaries from 10 miles
northeast of Moab, Utah, to Lake Powell.
Many populations of this species disappeared
with the impounding of water in Lake Pow-
ell.
Atriplex ganettii is a very fragile species
and would probably be facing extinction
were it not for the protection afforded by its
inaccessibility in the steep canyons and nar-
row gorges along the stretch of the river
where it grows.
The third diploid form is restricted to the
Little Sahara sand dunes in central Utah. It is
strikingly different from other A. canescens
in its gigas habitat. Its growth rate is nearly
twice that of the tetraploid forms that grow
nearby (Stutz and Melby 1968). This rare
form is becoming increasingly threatened.
The sand dune retreat has apparently pre-
served it to date by excluding herbivores that
have difficulty in walking on the dunes. Re-
cent development of recreational facilities on
the dunes by the BLM as a resort for dune
buggy enthusiasts may spell its doom. Many
plants are damaged directly by dune buggies;
others are destroyed by people. Because this
is almost the only woody plant available in
this area, it is sometimes used as fuel. It is
also highly palatable to livestock and has
been harvested to feed horses. Other uses in-
clude mattresses for sleeping bags and make-
shift windshelters. The handsome fruiting
stalks are often gathered for home decora-
tion. During the annual spring dune buggy
racing events, thousands of people swarm
over these dunes. Even the games they play
take a toll.
Although requests have been made to pro-
tect this fragile population by restricting ve-
hicle use to a small area, it is apparently go-
ing to be difficult to accomplish. This is a
rare and endangered form which, although
identified by a very vocal sponsor, has still
failed to make the roster.
B. Tlie tetraploids
Although collectively the tetraploids are
abundant and widespread, numerous local-
ized small populations are genetically
imique. Many of these are obviously of signif-
icant biological value. Several deserve and
need protection; others appear capable of
holding their own.
Although some of the variation between
tetraploid forms may reflect separate poly-
ploid origins, most variations occur as prod-
ucts of interspecific hybridization. The fol-
lowing three examples are among the most
common.
(1) Atriplex canescens X confertifolia.
Hybrids between these very different spe-
cies have been previously reported by Plum-
mer et al. (1957), Plummer and Drobnick
(1966), and Hanson (1969).
The first one I found was in Elko County,
Nevada, 10 miles west of Wendover. It was
in an area which later became the median
between the lanes of a freeway and was
124
Great Basin Naturalist Memoirs
No. 3
therefore destroyed. From seeds harvested
from it, however, 17 seedHngs were obtained
which are now growing in the BYU nursery.
All of these plants appear very much alike in
habit and leaf characteristics but are dis-
tinctively different from either parent and
from all other species oi Atriplex. Differences
in fruit characteristic are apparently due to
only a few genes which permit clear segrega-
tion in this small population. Other charac-
teristics, such as habit, spininess, and leaf
characters, do not conspicuously segregate,
suggesting a more complex genetic control.
Surprisingly, the males show regular meiosis,
which implies that most of the differences ac-
quired by these two very distinct species
have come from gene mutations unaccompa-
nied by gross chromosomal aberrations.
A large number of hybrids from this par-
entage have now been collected and progeny
from them assembled in a common garden.
From these, it appears that at least some of
the natural variation present in both parental
species has come from introgression from
these hybrids. Near Honey Lake, California,
a small population of A. canescens is ob-
viously heavily introgressed with A. confer-
tifolia genes. Near Garrison, Utah, a popu-
lation of A. confertifolia appears to have
received genes from A. canescens.
Should these hybrids and introgressed pop-
ulations be included on the rare and endan-
gered roster? Individually, they appear to be
fully qualified. They are highly important as
sources for new adaptive combinations. Some
possibly might represent the beginnings of
valuable species if they are spared. Will they
need names to make the preferred list?
(2) Atriplex canescens X A. cuneata.
Although A. canescens is a large woody
plant and A. cuneata is a low-statured her-
baceous perennial, hybrids between them are
common. They have been reported by Plum-
mer et al. (1957), Plummer and Drobnick
(1966), and Hanson (1969). Stavast (unpubl.
no.) reported an extensive population of
hybrid segregants west of Hanksville, Sevier
Coimty, Utah.
From natural hybrids, segregating seed-
lings have been grown to maturity in the
BYU nursery. While most of them are more
like A. canescens than like A. cuneata in
habit and fruit characteristics, cuneata in-
fluence is unmistakable.
From observations of these garden-grown
segregants, it has since been possible to iden-
tify several distinct introgressed populations
in nature. One particularly striking form is
becoming established near Ferron, Emery
County, Utah, as a low-statured form, with
small fruits and a capacity for growing with
and favorably competing with, Ceratoides la-
nata and Xanthocephalum sarothrae, which
neither parent can do. It now occupies only
about 40 acres so is still rare and, of course,
endangered. It may be the beginning of a
very valuable addition to our rangelands if
we can preserve it.
Other novel adaptive combinations from
this same parentage are likely also forthcom-
ing if the source is protected.
(3) Atriplex canescens X A. gardneri
Many years ago A. Nelson reported hy-
brids between A. canescens and A. gardneri
and named them A. aptera (Nelson 1904).
Such hybrids are still common west of La-
ramie, Wyoming, and elsewhere where these
two species meet. They appear to have given
rise to a series of very successful derivatives
which now occupy the banks of most of the
tributaries of the Missouri River in Montana,
southern Alberta, southern Saskatchewan,
North and South Dakota, and northwestern
Nebraska. It was a specimen of this form
which was collected by Lewis and Clarke in
1804 north of Chamberlain, South Dakota,
and to which the name A. canescens was as-
signed. In most places it is low growing and
shows vigorous root-sprouting, but some are
quite woody. Some have broad wings on the
fruiting bracts; some show only small traces
of wings. Apparently segregation is still going
on as unique combinations find habitats in
which they are competitive.
Collectively these hybrid products are not
at all rare or endangered, but local unique
populations certainly are. Should tliey re-
ceive protection?
Several other interspecific hybrids in-
volving tetraploid A. canescens from which
segregating progeny are sometimes abundant
and stabilized, sometimes rare and variable,
have been found in western North America.
Some have already yielded new adaptive in-
1979
The Endangered Species: A Symposium
125
cipient species; others may yet do so from
the rich common gene pool. Some of them
will obviously require protection if they are
to become established. Others already appear
to be sufficiently established to be able to
continue even with human assaults.
C. TJie hexaploids
Within many tetraploid populations of A.
canescens, occasional hexaploid plants have
been found. They are apparently contin-
uously and sporadically produced from unre-
duced gametes. For the most part, they have
not become established as separate adaptive
derivatives. This is probably due primarily to
the high improbability of two such hexaploid
plants being produced simultaneously in suf-
ficiently close proximity to each other to in-
terbreed, and also, even if they did, it would
be imusual for their offspring to be an im-
provement over the parental forms.
Even so, a few exceptional, small, localized
hexaploid populations have been found in
isolated pockets. In the White Sands National
Monument, New Mexico, some very promis-
ing hexaploids have become established in lo-
calized colonies fairly close to the more
abundant tetraploid form. In the BYU nur-
sery, they show a shorter, more compact
habit than the tetraploids and may have attri-
butes which would be superior in particular
range conditions. Because they are still few
in number and sporadic in distribution, they
could profitably use protection, but under
current policy they appear to have little
chance of receiving it though their existence
under circumstances that severely hinder
their establishment suggests they may have
great potential for success once they get
started.
The ephemeral hexaploids that appear as
single plants in tetraploid populations, al-
though rare and of course endangered, prob-
ably do not merit legislated protection, sim-
ply because they cannot demonstrate
particular values until removed from their
tetraploid neighbors and manipulated by
plant breeders. Their potential may be high,
but protecting them in their ephemeral in-
fancy is probably not warranted, although
they apparently meet all current prerequi-
sites except for having designated names.
In contrast to rare, ephemeral, and local-
ized small pockets of autohexapaloids, there
are several allohexaploids that appear to hold
great promise as new additions to the desert
ranges. One of these is already abundant in
the north-south-oriented valleys of Nevada. It
appears to have been derived from the par-
entage 4N A. canescens X 2N A. falcata.
Apparently doubling of the chromosomes in
the triploid F^ hybrid gave rise to a remark-
ably well-adapted new taxon. It is a short-
statured form that often flowers during its
first year of growth. Because there are
marked differences between different popu-
lations, they have apparently arisen repeat-
edly at different places. Although, collec-
tively, this hexaploid is well established,
successful, and apparently capable of with-
standing intensified grazing pressures, some
of the individual component populations are
genetically distinct and apparently suffi-
ciently rare to be endangered. Should we at-
tempt to protect these new arrivals during
their fragile infancy, or shall we settle for the
already acquired forms which are performing
at least satisfactorily?
Another hexaploid fourwing derivative
that appears to offer unique and exciting
promise as a new adaptive taxon, has appar-
ently come from the parentage 4N A. canes-
cens X 6N A. tridentata. This interesting
form occupies only about 80 acres east of
Grantsville, Tooele County, Utah. It is up-
right and woody like typical nearby tetra-
ploid A. canescens plants, but it has soft-tex-
tured furfuraceous leaves and late-maturing
flowers and fruits like A. tridentata. It also
grows on heavy clay soils as does A. triden-
tata.
This small population appears to be re-
markably adapted to this valley and is appar-
ently spreading. Currently almost entirely
within a military reserve, it is already receiv-
ing needed protection during its infancy.
However, if that protection were removed,
the entire population could very quickly be
lost. Indeed, were it not for the presence of
the reserve, it may have never survived
beyond its birth.
A hexaploid A. tridentata-\ike derivative
from this same parentage has apparently
come into existence only during the past dec-
ade. It is still confined to the roadsides along
a 30-mile stretch of freeway between Salt
126
Great Basin Naturalist Memoirs
No. 3
Lake City and Wendover. Because the free-
way itself is only about a decade old, the new
adaptive derivative must also be no older
than that. In the center of the population, A.
tridentata and A. canescens are sympatric.
Hybrids and hybrid products, as well as the
new stabilized segregant, are all present. In
the summer of 1977, an actual count was
made of the plants of this new form. On the
roadsides of the lanes leading westward into
Wendover, 17,600 plants were counted. As-
suming approximately the same number on
the roadside of the lanes leading eastward to-
ward Salt Lake City, the total population
consists of only about 35,000 plants. It is still
rare, but as long as the highway is there there
is little threat to its continuation. This por-
tion of the freeway is mostly across empty
salt flats, so grazing and other biological
pressures are essentially absent. Conceivably,
these robust, unique plants may be pre-
adapted for occupying areas other than the
side of the freeway, in which case they may
one day find an escape from this restricted is-
land, in any case, legislated protection is
probably meaningless, despite their rarity
and high intrinsic aesthetic and scientific val-
D. The twelve-ploids
Atriplex canescens var. laciniata Parish is
distinguished by fruit bracts that are thin and
lacy. It is common sporadically in much of
the Mojave Desert, with extensive popu-
lations near Barstow, California, and around
the Salton Sea. It is apparently an allopoly-
ploid derived from the hybrid A. canescens
X A. pohjcapra.
Although this extraordinary species is now
well established and apparently in no need of
protection, it hybridizes freely with both A.
canescens and A. pohjcarpa, yielding numer-
ous novel progeny, each one of which is rare.
Although most are aneuploids, some of them
may be preadapted for habitats yet uninha-
bited.
Parallel examples can be drawn from other
groups of saltbushes and probably from many
other desert plants. Rare taxa and endan-
gered taxa are commonplace in these rapidly
changing environments. The problem then is
not one of finding them or defining them.
but, rather, understanding them. Not until we
know their biology and their genealogy can
sound decisions be made concerning their
management. Large, genetically uniform
populations may be, biologically, much more
endangered than smaller but heterogenous
populations. Genetically they are certainly
more rare.
In terms of management then, it is far
more important to identify rare and endan-
gered genotypes than rare species. In some
instances species having abundant genetic
variation and few individuals may be much
less endangered than species having limited
genetic variation, albeit many individuals.
Conclusion
It appears clear, therefore, that the mean-
ing of "rare and endangered" must extend
beyond mere head counting. Abundant indi-
viduals may not always mean abundant and
therefore secure genotypes, and vice versa:
species represented by only a few individuals
may be so rich genetically that their continu-
ance, under almost any normal environmen-
tal assault, is essentially certain. Equally im-
portant, in view of the cost to ecosystems, to
human society, and to other contemporary
organisms, some rare forms may not warrant
preservation at all. A sterile, weak poly-
haploid derivative with essentially no poten-
tial for amounting to anything of value can-
not justify protective measures merely
because it is rare and endangered. A dinosaur
pet, as fun as it might be to have, would be
prohibitively costly just to feed— let alone to
house and to exercise. On the other hand,
new exciting infant forms with rich potential
for high aesthetic, biological, economic, or
academic values should be encouraged and
their establishment and growth accelerated.
To tell which of the rare forms are coming
into existence and which are going out re-
quires intimate knowledge of their biology
and genealogy. Simply enumerating named
taxa which are rare is not enough. If we are
going to meddle in the evolutionary process,
let us do it intelligently. Otherwise it would
be better that we do not meddle at all.
1979
The Endangered Species: A Symposium
127
Questions for Dr. Stutz
Q. Howard, how are you going to choose wliicli ones
voii are going to save?
A. I would expect that vakie judgements will be used
just like we use them in creating priorities in every
decision we make in life. We categorize them. So I
think that if I were given an array of choices I could
make that decision on the basis of relative value. But
it's not going to be a simple recipe. It's going to be
based on intimate knowledge of the species being
considered.
Q. But you mean they are going to be entirely based on
man?
A. In the absence of that, then we would have to do as
has been suggested by several, including Dr. Steb-
bins and Dr. White. We simply maintain the ecosys-
tems, refrain from imposing our personal prefer-
ences, and let natural selection make the choice.
Then we're removed from that dilemma. I think, in
many cases, ultimately that's what has to happen.
It's obvious we cannot put everything into a wilder-
ness protected area, but we need to have preserves.
We need to provide opportunities where the evolu-
tionary process can proceed without our inter-
vention. On the other hand, there are situations
where we must evaluate. We will have to decide
whether to plow that field or to put in that hydro-
electric plant. When it is necessary to impose hu-
man decisions, then we must also impose human val-
ue judgements. At that point we need all of the
biological information we can possibly get to make
those decisions. Thev must not come simply from a
knowledge of numbers of individuals alone.
Dr. Stebbins: First I think this is a fascinating topic, but
I'd like to bring this whole question of preserving or
not preserving into the context of what you said
about intelligent meddling with evolution. I think in
this case, if we're going to understand evolution, we
want to make it go. After all, the engineer doesn't
just look at what electric motors did in the past, he
makes new ones. Now in the case of the examples
you have in both your Purshia-Cowania and your Af-
riplex, it impresses me that these obviously recent
populations have not yet fitted in to any jDarticular
ecological niche. The way to save them, in my opin-
ion, is to gather large numbers of seeds and meddle
just a little bit by finding out just what kind of eco-
logical niches they prefer. Those that aren't likely to
be disturbed will be happy homes for these things
and will then lead on to something new and still dif-
ferent. I'm particularly interested in this stabilized
Purshia colony in that connection. You should go up
there and ask permission from that rancher to get all
the seeds you can and grow them somewhere along
the margin there and just see if you can't find a
place where it will be more than just a puny little
population. In the case of your Wendover freeway,
heavens knows there are miles and miles and miles
of freeway that have nothing but Salsola kali, for in-
stance. Wouldn't it be nicer to have this thing rather
than Russian thistle or halogeton?
A. The answer is in the affirmative. I'm glad vou
brought that into perspective. Dr. Stebbins, because
we have another great opportunity before us today
which needs to be exploited. That is the sudden
availability of new environments provided by mining
operations in which we can do this very thing. Dr.
Frischknecht of the Forest Service Shrub Laboratory
is working on preparing plants which will be able to
tolerate oil-shale refuse dumps. Also with strip min-
ing, there are brand new islands made available and
new areas in which we can do just exactly as Dr.
Stebbins suggested. We can introduce gene pools
into these new arenas and watch them evolve. We
can monitor the evolutionary changes and we can
get a record of evolutionary dynamics like we've
never been able to before. We need to cooperate
with industry and use their by-products to help us
learn more about succession and evolution.
Dr. Stebbins: Let me just mention this. I think it's nov-
el, but I don't know how many of you know about it.
If you have read the books of the marvelous scien-
tific philosopher, Rene Dubois, he has made the
comment that we Americans are too wilderness ori-
ented becau.se we, or our ancestors, were brought up
in or near pioneer habitats and wilderness, whereas
Dr. Dubois was brought up in the vicinity of Paris.
He knows country as cultivated land, as well-mani-
cured forests, and sees the beauty in that. Isn't there
some justification in our thinking in terms of produc-
ing a pleasing landscape of human manufacture from
many of the areas which are just junk now and, at
the same time, of course, preserving the wilderness?
Dr. Deacon: A couple of comments, one fairly specific.
It is possible to include unnamed entities. I think
that's one of the criteria that need not be met. The
listing process that I've been involved with has
something like over 10 percent unnamed taxa.
Dr. Stutz: How are they listed? With a number?
Dr. Deacon: They're simply referred to as a subspecies
with a common name that's distinctive or unique to
that group. In other words, what is necessary is to
realize that it is unnecessary to actually go through
the process of naming.
Dr. Stutz: This entity, for example of Purshia-Cowania.
I suppose we'll need a handle before it can even get
on the roster. That was my only point.
Dr. Deacon: Not the formal scientific name. But more
serious than that, in my view, was the illustration of
the coyote/rabbit: if you .save one, you're likely to
save the other. It's the same sort of illustration that
Congressman McKay used with respect to the Colo-
rado squawfish eating the humpback chub. The
point is, if you save an evolving ecosystem you save
all parts of it. Just because you kill an individual
doesn't mean you kill the species so that the evolu-
tion of predator/prey is what nuist be preserved. I
would hope you might reconsider using that illustra-
tion.
Dr. Stutz: I already have.
Dr. Deacon: The other point I would like to make in
that respect is that certainly the consideration of
value, which is the main point of your talk, is really
the most difficult thing we have to deal with here,
and when you come to the process of involving eco-
nomic value, it looks like one of the most fniitfid
possibilities for consideration. The discussion pre.s-
128
Great Basin Naturalist Memoirs
No. 3
ented so nicely to us yesterday by Dr. Spencer is
perhaps one of the most optimistic I've heard pres-
ented here from the standpoint of the pressure al-
ready in existence. I think he represented to us the
changing social values that are in fact forcing us into
the changes necessary for us to establish "a world-
wide sensible economic system."
Mr. Clement: This is a fascinating evolution in refining
our expression of what we're concerned about, and
let me add one more fact: distinguishing between
economic and fiscal valuation. Most of what we call
economic is fiscal, private concern about economic
return, and when you come to valuing in a broad
sense, all values are economic, because we're dealing
with scarce resources, whether they're material, aes-
thetic, or spiritual.
Literature Cited
CoTTAM, W. P., J. M. Tucker, and R. Drobnick. 1959.
Some clues to Great Basin postpluvial climates
provided by oak distributions. Ecology
50:361-377.
Drobnick, R., and A. P. Plummer. 1966. Progress in
browse hybridization in Utah. Proc. Conf. West.
State Game and Fish Comm. 46:203-211.
Hansen, C. A. 1962. Perennial Atriplex of Utah and
northern deserts. Unpublished thesis. Brigham
Young University, Provo, Utah.
Plummer, A. P., R. L. Jensen, and H. D. Stapely. 1957.
Job completion report for game forage project.
Inform. Bull. 31. Utah State Dept. of Fish and
Game.
Pyrah, G. L. 1964. Cytogenetic studies of Cercocarpus
in Utah. Unpublished thesis. Brigham Young Uni-
versity, Provo, Utah.
Stutz, H. C. 1978. Explosive evolution of perennial Afn-
plex in western America. Great Basin Nat. Mem.
2:161-168.
Stutz, H. C, J. M. Melby, and G. K. Livingston. 1975.
Evolutionary studies of Atriplex: a relic gigas
diploid population oi Atriplex canescens. Amer. J.
Bot. 62:2.36-245.
Stutz, H. C, and L. K. Thomas. 1964. Hybridization
and introgression in Cowania and Purshia. Evolu-
tion 18:18.3-195.
SOME REPRODUCTIVE AND LIFE HISTORY CHARACTERISTICS
OF RARE PLANTS AND IMPLICATIONS OF MANAGEMENT
K. T. Harper'
.\bstract.— .\nalvsis of the vascular floras of Utah, Colorado, and California suggest that a syndrome of life form
and reproductive characteristics separates rare and common species. Woody plants are heavily underrepresented,
and herbs are overrepresented on the official lists of endangered and/or threatened plants of the floras considered.
Few of the rare species are descended from wind-pollinated ancestors, but instead are derived from insect-pollinated
stock. Theorv suggests that many of the rare taxa will ultimately be .shown to be self-pollinated. The date show a
tendency for rare species to be better represented among taxa having bilaterrally symmetrical as opposed to radially
symmetrical flowers. In aggregate, the results suggest that most rare taxa are equipped for rapid exploitation of
small, unusual habitats. Because many rare taxa appear to be dependent on insects for reproduction, their survival
depends not only on appropriate physical habitat but also on healthy pollinator populations. Reproduction of out-
crossed taxa will be handicapped by road dust and other sources of atmospheric particulate which might foul stig-
matic surfaces. Self-pollinated taxa may have little generic variability and thus be especially sensitive to environmen-
tal modifications. Because most rare taxa are dicotyledonous herbs, herbicides such as 2, 4-D which have been
widelv used in vegetation management for control of broadleafed plants can be expected to have highly deleterious
effects on populations of rare species in the target area.
The goal of management in any discipline
is control of the components of the system
mider consideration. The components of any
system can be controlled only if their charac-
teristics are imderstood. Once the critical
characteristics and their dynamics through
time are known, control strategies can be for-
mulated and tested.
Currently we know a great deal about
which plant species are so uncommon that
their existence could be endangered by even
moderate natural or manmade changes in the
environment. We know less about the size
and distribution of the populations of most
rare species. Even less is known about the
habitat requirements of the individual rare
species. But perhaps our greatest ignorance
concerning rare taxa relates to the specifics
of their life history and reproductive biology.
Before such taxa can be successfully man-
aged, managers must understand the life
cycle, longevity, and reproductive habits of
each.
In this paper, I will examine various life
form, longevity, and reproductive characters
of endangered and /or threatened plant spe-
cies of three states of the western United
States: California, Utah, and Colorado. The
incidence of a given characteristic among
species listed as endangered or threatened in
a particular state (U.S. Department of Interi-
or 1975) will be compared to the incidence
of that character in the entire seed plant
flora of that state. By the use of appropriate
statistical tests, characteristics that are over-
or underrepresented among rare taxa can be
identified, provided the incidence of each
character is known among both rare taxa and
the full flora of the state.
Methods
The basic data for this paper have been
drawn from Munz and Keek's (1973) flora of
California, a Soil Conversation Service
Checklist of Utah plants (no date given), Har-
rington's (1964) flora of Colorado, and the
U.S. Department of Interior's (1975) initial
listing of endangered and threatened plant
species. Characteristics of the individual taxa
have been drawn from species descriptions in
the floras, examination of herbarium mate-
rial, and personal experience. Some taxa in
all states could not be characterized ade-
quately and were thus omitted from consid-
'Department of Botany and Range Science. Brigham Young University, Provo, Utah
129
130
Great Basin Naturalist Memoirs
No. 3
eration (this deficiency was particularly
serious for the California flora).
Suffrutescent (woody rooted herbs) taxa
are treated as perennial herbs. Annual and
biennial taxa were combined for the purposes
of this paper. Mode of pollination (wind, wa-
ter, or animal) was inferred for each taxon
from floral structure, degree of exertion of
stamens and stigmas, and published reports.
The accessibility of the pollen and/or nectar
to the average animal visiting the flower was
deduced from floral structure. Flowers were
considered to have restricted access to ani-
mals if they possessed any of the following
characteristics: (1) petals, sepals, or calyx
tube fused into a long (over 3 mm) tube of
small diameter (as in some Gilias, Oeno-
theras, or Cirsiums), (2) nectaries positioned
in tubes that extend away from the reproduc-
tion organs (as in Delphinium or Aguilegia),
and (3) separate sepals and petals that are so
positioned as to stand rigidly erect forming a
narrow, false tube around the reproductive
organs (as in Erysimum, Streptanthus, or
Vicia). Not all sympetalous taxa were consid-
ered to have restricted access flowers. For in-
stance, some Campanula, Valeriana, and
Kabnia species were classified as being open
and freely accessible to pollinators. Although
composite flowers that include both ray and
disk flowers could be considered to include
both radially and bilaterally symmetrical
components, I have classified such flowers as
radially symmetrical. Composite flowers con-
sisting of ray flowers alone have also been
categorized as radially symmetrical in this
study.
For the purposes of this paper, I have as-
sumed that characteristics that are over-
represented among rare taxa (in comparison
to the flora from which they have emerged)
impart some survival advantage to the rare
entity. Conversely, I assume that character-
istics that are underrepresented put rare taxa
at a survival disadvantage. It will be recog-
nized that the foregoing assumptions are
based upon yet another assumption, namely
that most taxa that are designated as endan-
gered or threatened are relatively recent in
origin. This latter assumption imphes that the
rare taxon is commonly possessed of a suite of
characteristics that permit it to be successful
in spite of small populations and restricted
habitat. I also recognize that the foregoing
assumptions reveal still another assumption
inherent in the analyses presented here: if
characters that are overrepresented among
rare taxa are viewed as enhancing their
chances of survival, it must be assumed that
ancient taxa that are not well suited to mod-
ern conditions and are thus in a state of pop-
ulation decline are uncommon entities on the
lists of endangered and threatened species.
This point will be considered further in the
Discussion section.
In the analyses which follow, the chi-
square statistic is used to determine whether
a characteristic is over- or underrepresented
among the taxa listed as endangered or
threatened: the incidence of that character in
the regional flora is used as the basis for com-
parison. In such analyses, the individual spe-
cies become the statistical observations or
replications in the compartments of the 2 X
2 contingency tables. Relationships were de-
clared statistically significant only when the
probability value for the relationship was .05
or less.
Results
Characteristics of Regional Floras
Five regional floras have been analyzed in
connection with this study (Table 1). Each
flora has been analyzed to give the incidence
of some or all of the following character-
istics: percentage of the taxa that are woody
(shrubs or trees), percentage that are wind
pollinated, percentage that are short-lived
(annual or biennial), percentage that have
flowers that do not restrict animal access to
pollen and/ or nectar, and percentage that
are radially symmetrical. The five floras are
surprisingly similar in respect to the fore-
going characteristics. The most different flora
in respect to the characteristics considered is
that designated as Wasatch Prevalent Spe-
cies. As the name implies, that list ignores
species that were sampled infrequently (Os-
tler and Harper 1978). An emphasis on the
more common species of a region seems to
unduly emphasize woody and wind-polli-
nated species: annuals appear to be under-
represented on the Wasatch Prevalent Spe-
cies list. Whether the underrepresentation of
1979
The Endangered Species: A Symposium
131
annuals is attributable to the fact that the
Wasatch Prevalent list ignores desert habitats
or to some other cause is unknown.
The date (Table 1) demonstrate that
woody species constitute between 10 and 15
percent of the state floras considered. In
areas that are primarily desert (such as the
Kaiparowits region of Utah), woody taxa may
contribute almost 25 percent of the species in
the flora. Wind-pollinated species contribute
from 19 to 26 percent of the species in the
regional floras studied. Short-lived species
(annuals or biennials) furnish from 22 to 31
percent of all species in the regional floras
imder consideration. Animal-polluted flowers
dominate all of the floras considered. In the
Colorado flora and the two subsamples of the
Utah flora, only from 32 to 41 percent of the
zoophilous taxa have flowers that are fully
opened (nectar and /or pollen readily reached
by most animal visitors). Most of the zoo-
philous taxa in the Colorado and Utah floras
are radially symmetrical (79 to 85 percent of
the species).
The characteristics of the regional floras
will serve as the basis against which charac-
teristics of the endangered and threatened
species of those floras will be compared. In
the case of the Wasatch Prevalent Species
list, no endangered or threatened species are
included. Consequently, characteristics of
species from the bottom third of the com-
monness gradient formed by arranging the
prevalent species in order of decreasing aver-
age frequency will be compared with charac-
teristics of those species which appear on the
top third of the commonness gradient. Hope-
fully, such an analysis will reveal something
about characteristics that enhance the survi-
val of less common species.
Size and Longevity of Rare Taxa
In four of the five floras examined, woody
plants are underrepresented among the en-
dangered and threatened taxa (Table 2). The
nonconforming flora is that of California:
there woody plants are more common among
endangered taxa than one would expect con-
sidering the number of woody taxa in the
state flora, but the departure from random
expectations is not statistically significant. Al-
though most of the endangered woody spe-
cies in California belong to three rapidly
evolving genera {Arctostaphylos, Ceanothus
and Eriogomim), a number of the taxa appear
to be old entities that are survivors of ancient
groups that are well adapted to only a few of
the modern environments of the state. Spe-
cies representative of apparently old, declin-
ing lineages include the following: Cupressus
goveniana var. abramsiana, Juglans hindsii,
Lavatera assiirgentiflora, Lyonothamnus
floribundus, Fremontodendron decumbens
Table 1. Number of species studied and characteristics of the floras considered. Floristic data sources appear at
the bottom of the table. Blanks occur in the table where specific analyses have not been made.
Flora
Wasatch
Kaiparowits'
Prevalent'
Characteristic
California'
Utah^
Colorado'
(Utah)
Species (Utah)
Size of flora
Percent woodv species
5,489
14.0
3,507
10.1
2,7.35
11.0
848
22.1
244
21.3
Percent wind pollinated
species
18.9
18.5
25.9
24.0
.33.2
Percent annual or biennial
species
Percent unrestricted access
flowers (Zoophilous only)
Percent radially
30.7
21.9
22.7
40.4
25.1
.32.0
12.3
39.9
symmetrical flowers
(Zoophilous only)
-
-
80.2
79.0
84.7
'Munz and Keck (1973)
•Soil Conservation Service (no date given)
'Harrington (1964)
'Welsh et al. (1978)
•Ostler and Harper (1978)
132
Great Basin Naturalist Memoirs
No. 3
and F. mexicanum. The floras of Utah and
Colorado appear not to have a significant
representation of such ancient, woody taxa.
The weight of the evidences seems in favor
of the hypothesis that larger (woody) species
are underrepresented among rare species.
Three reasons may be suggested for the un-
derrepresentation of woody plants among
rare taxa: (1) large size limits the number of
individuals that can occupy any given area,
(2) slower maturation rates are accompanied
by lower rates of population growth, all oth-
er things being equal, and (3) long life and
low reproduction rates impede the rate at
which genotypes can be attuned to peculiar
environments. The affect of organismal size
on individuals per unit area is self-evident.
The profound influence of age at first repro-
dviction on intrinsic rate of increase of a pop-
ulation was demonstrated over a quarter of a
century ago by Lamont C. Cole (1954 and
Fig. 1). Unquestionably, the average age at
onset of reproduction is older for woody
plants than for herbs. Thus organismal size
and age at first reproduction can be expected
to combine to depress the population size of
woody taxa in the early history of their exist-
ence. Theory strongly supports the concept
that extinction rate is inversely correlated
with population size and intrinsic rate of re-
production (Pielou 1969:17). Theorists con-
clude that most extinctions occur during the
initial phase of population growth (Ricklefs
1979:649). Unfortulately, that is the period
when slow-maturing organisms such as
woody plants are at a particular disadvantage
in terms of reproduction rate and population
size. The chances of extinction for woody
species is further enhanced by a slow rate of
genetic fine-tuning to unique environments.
Small, faster-reproducing (because of earlier
maturation), and short-lived herbaceous taxa
are almost certain to genetically adapt to
new environments faster than woody taxa.
Given the advantages of small size and
early reproduction, one might have expected
Table 2. The observed and expected occurrences of woody taxa among rare species of five floras. Expected values
are based on the occurrence of woody taxa in the regional floras. A sample 2x2 contingency table appears at the
bottom of this table. Expected values in the contingency table appear in parentheses.
Flora
Characteristic
California
Utah
Colorado
Wasatch
Kaiparowits Prevalent
(Utah) Species (Utah)
No. of endangered and
threatened species
considered 234'
No. of woody species
observed 40
No. of woody species
expected 33.0
Chi-square summation for
the relationship 1.8
Significance of relationship NS
157
5
15.1
7.8
49
1
5.3
4.0
44
5
9.5
2.9
NS
85^
14
22.9
9.7
Life form
Species group
Herbs
Woody
Total Taxa
California endangered
California flora
194
(201)
4,721
(4,714)
40
(33)
768
(775)
234
5,489
4,915
808
5,723
'Endangered species only considered
'No. endangered or threatened species included: species from the bottom third of the commonness gradient used instead
NS— not statistically significant
■-statistically significant at the .05 but not the .01 level
"—statistically significant at the .01 level
1979
The Endangered Species: A Symposiui
133
annual and biennial plants to be significantly
overrepresented among the rare taxa. In only
the California flora, however, were the short-
lived taxa overrepresented, and even there
the relationship fell far short of statistical sig-
nificance (Table 3). In both Utah and Colo-
rado, annuals and biennials were significantly
underrepresented. Two possible reasons are
offered for the results observed: (1) short life
requires that a genotype by highly pre-
adapted to the environment to be occupied,
and (2) the relationship may be a taxonomic
artifact because annual and biennial groups
appear not to have received the close tax-
onomic scrutiny that numerous perennial
herbaceous and wood plant groups have been
exposed to. In respect to reason 1, many
unique perennial herbs undoubtedly persist in
potentially exploitable environments for
many years before genetic recombinations
are generated that permit the taxon to suc-
cessfully colonize the site. Such extended pe-
riods of genetic "experimentation" would not
be possible for annual or biennial taxa: in
their case, the novel genotype must reach an
open niche and be sufficiently well adapted
to that environment to reproduce success-
fully in the first reproductive event. The
probability that the novel genotype will be
sufficiently preadapted to reproduce success-
fully in the potential niche during the first
reproductive event is apparently small.
2.4-.
2.2
2.0-
1.8-
c/)
O
.E 1.4-
H—
O 1.2 -
TO 10-1
.9 0.8.
</)
C
'^ 0.6-1
_C
0.4-
0.2-
b = litter size
n = no. reproductive events /O
15
Age of first reproduction
Fig. 1. The influence of age at first reproduction on rate of natural increa.se in population size. Note that delaying
reproduction for even one reproductive period (from period 1 to period 2) for taxa that produce many offspring per
reproductive event has a major effect on the intrinsic rate of poulation increase (about a 45 percent decline when b
= 10 and the females reproduce repeatedly). Many perennial herbs reproduce in the first year of life, but many
woody taxa delay reproduction for more than a decade. (Figure modified from Cole [1954]).
134
Great Basin Naturalist Memoirs
No. 3
Table 3. Observed and expected occurrences of short-lived (annual and biennial) taxa among rare species of five
floras. Expected values are based on the occurrence of short-lived taxa in the regional floras. Number of endangered
and threatened species remains as in Table 2.
Flora
Wasatch
Characteristic
California
Utah
Colorado
Kaiparowits
(Utah)
Prevalent
Species (Utah)
No. of short-lived taxa
observed
80
15
4
9
9
No. of short-lived taxa
expected
72.2
33.,5
11.0
11.0
10.2
Chi-square simimation for
the relationship
1.3
13.6
5.8
0.5
0.3
Significance of relationship
NS
oo
"
NS
NS
.NS-Not statistically significant
•-Statistically significant at the .
"-Statistically significant at the .
05 but not the .01 level
01 level
Pollination of Rare Species
In all floras considered here, wind-polli-
nated taxa are underrepresented among the
rare species. The relationship is significant at
the .01 level in four of the five floras consid-
ered (Table 4). As Ostler and Harper (1978)
have argued, wind pollination would be ex-
pected to be ineffective where species are
represented by few, widely scattered individ-
uals and where the species are small and
overtopped by larger plants. Because small
populations are a characteristic of all taxa
listed as threatened or endangered, wind pol-
lination would be expected to be a poor re-
productive strategy for them. In addition, the
threatened and endangered lists considered
consist primarily of herbaceous (and thus
small) organisms. Both Ostler and Harper
(1978) and Freeman et al. (1979) show that
wind pollination is heavily underrepresented
among herbs, apparently because they are
usually overtopped by woody vegetation that
restricts wind flow and hinders movement of
pollen to receptive stigmas.
Flower Symmetry
The data show a universal over-
representation of bilaterally symmetrical
flowers among the rare taxa of all floras
(Table 5). It must be noted, however, that the
values reported for California and Utah are
conservative estimates only; the actual in-
cidence of radially symmetrical flowers in
those areas has not been tabulated directly
Table 4. Observed and expected occurrences of wind-pollinated taxa among rare species of five floras. Expected
values are based on the occurrence of anemophilous taxa in the regional floras. Niunber of endangered and threat-
ened species remains as in Table 2.
Flora
Wasatch
Kaiparowits
Prevalent
Characteristic
California
Utah
Colorado
(Utah)
Species (Utah)
\o. of vvind-polliriated
taxa observed
21
1
4
6
19
No. of wind-pollinated taxa
expected
43.3
27.9
15.5
10.4
30.0
Chi-square summation for
the relationship
14.7
33.0
12.7
2.6
12.7
Significance of relationship
°°
oo
oo
NS
o.
NS— Not statistically significant
•-Statistically significant at the .05 but not the .01 level
••—Statistically significant at the .01 level
1979
The Endangered Species: A Symposium
135
(see Table 5 legend for assumptions for Cali-
fornia and Utah). If the assumptions are valid
for Utah, bilaterally symmetrical flowers are
significantly overrepresented among rare
plants there.
The overrepresentation of zygomorphic
flowered species in the Kaiparowits flora nar-
rowly misses significance at the .05 level. In
aggregate, the data suggest that floral zygo-
morphy conveys a reproductive advantage to
rare plants. That advantage perhaps lies in
the fact that zygomorphy forces pollinators
to approach flowers in a stereotyped way.
Under such conditions (i.e., predictable posi-
tioning of the pollinator in the flower), selec-
tin can operate to position stamens and
stigmas within the flower so as to enhance
the efficiency with which pollen is trans-
ferred from stamen to pollinator and from
pollinator to stigma. Zygomorphy may also
enhance the distinctiveness of flowers of dif-
ferent species and provide another cue to
compliment color, size, and odor as charac-
ters that permit pollinators to distinguish
flowers of one taxon from those of another.
One would expect the degree of flower
uniqueness to enhance fidelity between the
flower and animal pollinators and thus im-
prove reproductive success.
Restricted Access to Flowers
Taxa with flowers in which access to nec-
tar and/ or pollen rewards is restricted are
overrepresented among the rare species in
the three floras for which floral structure is
known, but the relationship is statistically sig-
nificant for the Wasatch Prevalent flora only
(Table 6). Using a conservative estimate of
the incidence of restricted access flowers in
the California and Utah flora (i.e., the rate
for Colorado from Table 1), contradictory re-
sults are obtained for California and Utah, al-
though the results are not statistically signifi-
cant for either flora. It seems reasonable to
assume that mechanical barriers that restrict
access of many insect taxa to nectar and pol-
len of a given plant species would encourage
fidelity between that plant and adapted pol-
linators, because adapted pollinators would
have greater assurance of a food reward at
each visit (i.e., many potential competitors
would be unable to harvest the floral re-
wards). The data lend only slight support to
the foregoing assumption, however, and it is
clear that mechanical hedges about floral re-
wards are much less useful adaptations to
rare plants than small size, early onset of re-
production, animal pollination, or bilateral
symmetry of the flower.
Table 5. Observed and expected occurrences of bilaterally symmetrical flowers among the animal-pollinated rare
species of five floras. Expected values are based on the occurrence of bilaterally symmetrical zoophilous flowers in
the regional floras except for California and Utah. Because the actual incidence of bilaterally symmetrical flowers is
unknown in the latter two floras, expected values are based on the conservative assumption that bilateral flowers
occur with a frequency in those floras equal to the frequency in the Kaiparowits flora (see Table 1).
Flora
Wasatch
Characteristic
California
Utah
Colorado
Kaiparowits
(Utah)
Prevalent
Species (Utah)
No. of threatened and
endangered species
considered
214'
156
45
38
66^
No. of liilaterally
symmetrical flowered
species observed
52
48
12
13
12
No. of bilaterally
symmetrical flowered
species expected
Chi-square summation for
the relationship
Significance of relationship
45.3
1.3
NS
33.5
8.4
9
1.3
NS
8.2
3.8
.10'
9.8
1.5
NS
'Endangered species only considered
'No. endangered or threatened species included: species from the bottom third of the commoness gradient used instead
'Statistically significant at the .10 but not at the .05 level
NS— Not statistically significant
•-Statistically significant at the .05 but not the .01 level
■"—Statistically significant at the .01 level
136
Great Basin Naturalist Memoirs
No. 3
Discussion
Results show that the CaHfomia flora be-
haves differently from that of Utah and Colo-
rado in respect to the frequency of both
woody and short-lived taxa. The over-
representation of rare woody taxa in Califor-
nia may be explained by the relatively high
incidence of apparently ancient woody taxa
there. Some of the ancient woody taxa of
California have been enumerated in the Re-
sults section of this paper. If there is a signifi-
cantly larger component of evolutionary old
taxa in California than in Utah or Colorado,
the divergent results reported in Table 2 for
California and the two interior states would
be expected. That is, in California ancient
woody species that are rare may be viewed
as taxa that were once more common but
have steadily lost habitat to more modern
species that are better adapted to current en-
vironments. If such is the case, the basic as-
sumption underlying this paper (i.e., charac-
teristics that are overrepresented among rare
taxa must enhance their chances of survival)
would not hold. Instead, characteristics con-
sidered to be selected against among recently
evolved taxa that have successfully eluded ex-
tinction may be overrepresented in older
floras that are now marginally adapted to
and declining in modern landscapes.
The slight overrepresentation of short-
lived taxa among endangered plants of Cali-
fornia (Table 3) is unique for the floras exam-
ined. It should also be noted that the Califor-
nia flora supports significantly more annual
plant species than any of the other floras con-
sidered (Table 1, Harper et al. 1978). It seems
likely that annual plants have been more in-
tensively studied in California than in the in-
terior states, but differences in taxonomic
treatment among the floras considered seem
inadequate to explain the differences noted in
Table 1 and in Harper et al. (1978, Table 3).
A suitable explanation for the apparent
greater success of annuals in California as op-
posed to Utah and Colorado is needed, but I
am unable to supply such an argument.
The fact that most of the rare taxa in the
floras considered are zoophilous or, at least,
derived from zoophilous stock (Table 4) sug-
gests the need for managers to use great care
when insect control programs are imple-
mented near populations of rare plants. Un-
less the taxa are self-pollinated or agamos-
permous (producing seed without
fertilization), decimation of pollinator popu-
lation would adversely affect reproductive
success of the plant species in at least the
year of treatment.
Unfortunately, the incidence of self-polli-
nation and agomospermy among threatened
and endangered species is almost totally un-
known. Consequently, all rare taxa should be
Table 6 Observed and expected occurrences of restricted access flowers among the animal-pollinated rare spe-
cies of five floras. Tfie number of endangered and threatened taxa remains as in Table 5. Expected values are based
on the incidence of flowers in which access to nectar and/or pollen is restricted in the regional floras except tor
California and Utah. Because the actual incidence of zoophilous species having restricted access to floral rewards is
unknown in the latter two floras, expected value is based on the conservative estimate that restricted access flowers
occur with a frequency in those floras that is equal to the frequency in the Colorado flora (see Table 1).
Flora
~
Wasatch
i
Characteristic
California
Utah
Colorado
Kaiparowits
(Utah)
Prevalent
Species (Utah)
No. of threatened or
endangered taxa
observed with restricted
flowers
119
99
29
26
45
No. of taxa expected to
have restricted flowers
127.1
9.3..3
26.9
25.5
39.1
Chi-square summation for
the relationship
Significance of relationship
1..3
NS
.9
NS
.4
NS
.04
NS
,5.6
NS— Not statistically significant
•-Statistically significant at the .05 but not the .01 level
1979
The Endangered Species: A Symposium
137
treated as obligate outcrossers until proven
otherwise. This point suggests that any man-
agement act that has the potential of dimin-
ishing pollen flow between separate individ-
uals of any rare taxon should be carefully
evaluated in terms of possible reproductive
impairment of that plant. Thus, construction
work or traffic over unpaved roads near pop-
ulations of either wind- or animal-pollinated
species should be carefully controlled or cur-
tailed completely, because dust can foul stig-
matic surfaces and essentially eliminate polli-
nation of obligate outcrossers.
A knowledge of the breeding system of all
rare taxa would markedly improve our ability
to make wise management decisions con-
cerning them. As noted above, outcrossing
taxa will necessitate more management re-
strictions than self-pollinated or agomos-
permous taxa. On the other hand, self-polli-
nated and agamospermous taxa may be far
less genetically diverse and hence more easily
disturbed by environmental alterations than
outbreeders. Furthermore, a knowledge of
the breeding systems of rare plants would
help phylogeneticists to better define the
probable origins of the taxa and geneticists to
estimate the likely amount of unique germ
plasm in given taxa (Baker 1961).
Circumstantial evidence suggest that many
of the threatened and endangered plant spe-
cies will be shown to be self-pollinated or
agamospermous. Such reproductive habits
would be expected to be selected for in rare
taxa for two reasons: (1) both habits would
tend to preserve unique gene combinations
that adapt rare plants to their habitat, and (2)
both reproductive habitats would permit soli-
tary individuals to successfully reproduce
(Grant 1971).
Finally, it is worthy of note for managers
that most of the rare taxa in all of the floras
considered here are dicotyledons. Thus, the
broad spectrum herbicides belonging to the
2, 4-D group (dichlorophenoxyacetic acid
and near relatives) that have proven so effec-
tive against broadleaved plants can be ex-
pected to be dangerous to most rare plants.
Herbicides of this group have been widely
used in land management programs in the
past for control of undesirable species. In the
future, threat to endangered species must be
added to the list of constraints that must be
considered when use of such herbicides is
considered.
Literature Cited
Baker, H. G. 1961. Rapid speciation in relation to
changes in the breeding systems of plants, pp.
881-885. In: Recent Advances in Botany, Univer-
sity of Toronto Press.
Cole, L. C. 1954. The population consequences of life
history phenomena. Quart. Rev. Biol. 29:10.3-1.37.
Freema.n, D. C, K. T. Harper, and W. K. Ostler.
1979. Ecology and plant dioecy in California and
Intermountain Western America. Oecologia 23.
In press.
Gr.\nt, V. 1971. Plant speciation. Columbia University
Press, New York.
Harper, K. T., D. C. Freeman, W. K. Ostler, and L.
G. Klikoff. 1978. The flora of Great Basin moun-
tain ranges: diversity, sources, and dispersal ecol-
ogy. Great Basin Naturalist Memoirs 2:81-103.
R\rrington, H. D. 1964. Manual of the plants of Colo-
rado. Sage Books, Denver, Colorado.
MuNz, P. A., and D. D. Keck. 1973. A California flora
(including the 1968 supplement). University of
California Press, Berkeley.
Ostler, VV. K., and K. T. Harper. 1978. Floral ecology
in relation to plant species diversity in the
Wasatch Mountains of Utah and Idaho. Ecology
59:848-861.
PiELOu, E. C. 1969. An introduction to mathematical
ecology. Wiley-Interscience, a division of John
Wiley & Sons, New York.
RicKLEFs, R. E. 1979. Ecology, 2d Ed. Chiron Press,
New York.
U.S. Department of Agriculture Soil Conservation
Service. No date given. List of scientific and
common plant names for Utah. 113 pp. (mim-
eographed).
U.S. Department of Interior, Fish and Wildlife
Service. 1975. Threatened or endangered fauna
and flora: review of status of vascular plants and
determination of critical habitat. Federal regis-
ter, vol. 40, no. 127, part V: 27,824-27,924.
Welsh, S. L., N. D. Atwood, and J. R. Murdock. 1978.
Kaiparowits flora. Great Basin Nat. .38:125-79.
THE IMPORTANCE OF BEES AND OTHER INSECT POLLINATORS
IN MAINTAINING FLORAL SPECIES COMPOSITION
V. J. Tepedino'
.\bstr.\ct.— Bees and other insect pollinators which are necessary for the successful reproduction of most species
of flowering plants, including agricultural crops, have been ignored by our preservation efforts. This is unfortunate
because bees, as low-fecundity organisms, are very susceptible to insecticides and populations are slow to recover
from perturbations. Many species of bees, particularly specialized species in the western United States and the trop-
ics, are vulnerable to extinction. With extinctions of specialized forms, generalized species, especially fugitives, are
expected to increase because of their ability to utilize a variety of resources and survive beyond the confines of
preserves. The possible effects of increased dominance by generalist pollinators on floral species composition is dis-
cussed.
Aside from being included in our objective
to preserve existent natural diversity (Ter-
borgh 1974), insect pollinators merit our
preservation efforts because some 67 percent
of extant flowering plants depend, to varying
extents, upon them for reproduction (Axelrod
1960). Indeed, "pollinators are an environ-
mental resource as critical to the long-term
survival of a (plant) population as are light,
moisture, etc." (Levin 1971). The adaptations
for the attraction and utilization of insects by
flowering plants for reproduction are impres-
sive. They include size, color, fragrance, nec-
tar, excess pollen, and nutrient contents, as
well as morphology, positioning, and devel-
opment of the floral parts (Percival 1965,
Baker and Hurd 1968, Faegri and van der Pijl
1971, Leppik 1972, Proctor and Yeo 1973).
In the absence of insects, most flowers as
they are produced today would be malad-
aptive and our flora would assume a different
aspect.
An example of a flora with few available
pollinators is that of the Galapagos Islands,
where only one species of bee and 19 species
of lepidoptera have been recorded (Linsley
1966). Where pollinators are in extremely
short supply it is disadvantageous to produce
large, attractive flowers. Instead, we expect
selection for wind pollination or autogamy
with a concomitant reduction in conspicuous
flowers (Rick 1966). In fact, there are few
brightly colored flowers in the Galapagos;
most are drab, and "endemics tend to have
reduced corollas" (Rick 1966). In pollination
tests with 18 species from seven families,
Rick found a high incidence of autogamy: 13
species self-pollinated automatically and one
was self-compatible. Results from four other
species were inconclusive. Linsley et al.
(1966) have speculated that successful in-
vasion of the islands may have been restrict-
ed to those plant species which are either
wind or self-pollinated or compatible with
available pollinators. Thus, the Galapagos
flora is probably less diverse than it might
have been had the pollinator diversity been
higher.
Bees are the most important of insect pol-
linators. Except for masarid wasps and a few
beetles, only bees depend exclusively upon
pollen and nectar for food throughout their
life cycle. Their coevolution with flowering
plants is manifest in the many morphological,
behavioral, and physiological adaptations
which make them more efficient at flower
utilization (Linsley 1958, Percival 1965, Bak-
er and Hurd 1968, Stephen et al. 1969,
Faegri and van der Pijl 1971, Proctor and
Yeo 1973).
In many cases the reciprocal adaptations
between particular bee and plant taxa have
'Department of Zoology and Physiology, University of Wyoming, Laramie, Wyoming. Current address: Bee Biology and .Systematics Laboratory. USDA-
SEA, AR, Utah State University, UMC 53, Logan. Utah 84322.
139
140
Great Basin Naturalist Memoirs
No. 3
become rather specialized. Within the genus
Penstemon, for example, species of the Series
Graciles are adapted for pollination by bees
of the genus Osmia (Crosswhite and Cross-
white 1966). Tropical orchids attract males of
particular species of bees of the genus Eu-
glossa by specific fragrances (Williams and
Dodson 1972). Pedicularis species are polli-
nated only by bumble bees (e.g., Macior
1977). Tlie pollen and nectar of the poisonous
range plant death camas (Zigadenus spp.),
though deadly to honeybees (Hitchcock
1959), are utilized by the oligolectic bee An-
drena astragali Viereck & Cockerell, which
also pollinates the plant (pers. obs.).
In addition to our native flora, many im-
portant agricultural crops, including cole
crops, orchard fruits and nuts, bushberries,
strawberries, some citrus fruits, sunflowers,
cucurbits, alfalfa, and red clover either re-
quire insect pollinators for seed set or set
more seed in their presence (Free 1970,
McGregor 1976). Although most crop polli-
nation is presently accomplished by honey-
bees, reports of native bees visiting flowers of
agricultural crops are common (Free 1970,
McGregor 1976), and endemics are undoubt-
edly responsible for some, as yet undeter-
mined, percentage of crop pollinations. Two
solitary species have replaced the honeybee
as the preferred pollinator of alfalfa in the
northwestern United States (Bohart 1972b),
and another species, Osmia lignaria Say,
shows considerable promise as a pollinator of
pome crops (P. F. Torchio, pers. comm.). Na-
tive bees will probably play an especially im-
portant role in the pollination of sunflowers,
a rapidly increasing native crop (F. D. Park-
er, pers. comm.). The preservation of exotic
bees will be important as we increase our ag-
ricultural acreages of introduced crops and
.seek to import pollinators that have coe-
volved with those crops. For example, the
primary pollinators of alfalfa in the north-
west, Megachile rotundata (Linnaeus), is an
exotic species native to Europe.
Of Grasshoppers and Bees
During the summer of 1978 a news release
in the Laramie Daily Boomerang announced
that a joint federal, state, and locally funded
insecticide spray program would be con-
ducted in northeastern Wyoming to control
grasshopper populations that had exceeded
economically safe levels. I paraphrase the last
segment of the news release: "Those who are
concerned about bee populations in the area
please note that all bees will be removed be-
fore spraying is conducted." This is quite an
impressive feat considering that Wyoming
has a minimum of 660 species (Lavigne and
Tepedino 1976), all but one of which evolved
with the native flora. The single exotic. Apis
mellifera, the honeybee, was, of course, the
species that was moved.
At the time insecticide was applied there
were probably between 25 and 50 species of
bees in the area. Unfortunately, no studies as-
sessing the affect of spraying were under-
taken, but other work has shown severe pol-
linator depletion following insecticide
application (Kevan 1975, Plowright et al.
1978, Robinson and Johansen 1978). Because
of their susceptibility to pesticides (Johansen
1977), wild bee populations were probably
decimated.
The rate of recovery of an animal popu-
lation whose numbers have been drastically
trimmed is positively correlated with the in-
trinsic rate of increase of that population
(May et al. 1974). Though females of many
insect species produce hundreds or thousands
of offspring, each of which receives little or
no parental care, bees have developed a con-
trasting strategy. Bees produce few offspring
and expend considerable effort to insure the
survival of each. Greenhouse studies at the
USDA Bee Biology and Systematics Labora-
tory with several solitary species, under con-
ditions of excess bloom and without natural
enemies, show that maximum fecundity aver-
ages 15-20 offspring per adult female. In the
natural environment, where bloom is only oc-
casionally superabundant and competitors,
predators, and parasites abound, fecundity
must be much lower. Because of their low fe-
cundity, bees recover slowly from bouts of in-
secticide spraving or other perturbations.
Plowright et al. (1978) estimated that three
to four years would be necessary for bumble
bee populations to return to prespray levels.
Such estimates assume cessation of spraying
and a continuously favorable environment.
Periodic spraying or long periods of weather
1979
The Endangered Species: A Symposium
141
unfavorable for flight would further slow re-
covery rates.
The low fecundity of bees has other ef-
fects. Smaller populations are more vulner-
able to local extinction by random events
(McArthur and Wilson 1967). If bee popu-
lations are periodically or consistently dis-
turbed, numbers will remain below carrying
capacity for extended periods, and popu-
lations will become more prone to random
extinction. The resistance to insecticides de-
veloped rapidly by many pest species is par-
tially due to the great genetic variability con-
tained in the prodigious numbers of offspring
produced by single females (Georhgiou
1972). Because of reduced fecundity, bees
may be less likely to develop resistance than
other insect species. Indeed, resistance to in-
secticides is unknown in bees.
The example of the bees that were not
moved (or even considered) illustrates our
philosophy of preservation. We have empha-
sized the preservation of species that are
"useful," closely related, or obvious to man.
We overlook the fimctionally important or-
ganisms that are frequently small and more
subtle in their actions. For example, insects,
the most influential of terrestrial animals
(man aside), whether judged by numbers of
species, individuals, or biomass, are repre-
sented by only eight threatened and endan-
gered species, all butterflies. Yet there are
over one million described insect species and
at least as many awaiting description. The
base of the trophic pyramid, plants, are rep-
resented by a mere 22 species. In com-
parison, 588 species of vertebrates appear on
the latest Threatened and Endangered Spe-
cies List (U.S. Department of the Interior
1977). A trend is evident even within the
class Vertebrata: 7.0 percent of all mammals
are threatened or endangered, 2.5 percent of
all birds, and 1.2 percent of all reptiles and
amphibians— but only 0.3 percent of all fish.
It is time that we attend to the preservation
of functionally important organisms without
backbones, many of which make vertebrate
existence possible.
The Rarity of Bees
As pointed out by Bohart (1972a), the ef-
fect of man on wild bees has been both posi-
tive and negative. Overall, however, bee
populations are probably in decline due to
habitat destruction and to our increasing de-
pendence on insecticides and herbicides. But
even this assessment is tenuous because of the
paucity of hard information. There are over
20,000 extant species of bees, and we know
almost nothing of all but a handful of them.
Our knowledge of tropical species is espe-
cially poor, but we can guess that with man's
rapid destruction of tropical habitats many
species will be lost. Even in the western
United States where bee diversity is very
high (Linsley 1958) we do not know how
many species, if any, have become extinct re-
cently or how many may be threatened. In-
deed, it is likely that the bee fauna of western
North America harbors many undescribed
species. For example, in two years of collec-
ting on shortgrass prairie in southeastern
Wyoming, I recorded over 200 species, 5-10
percent of which are new to science.
We do know from museum records that
many species are rarely collected. Lists of
such species could be compiled, but are these
species truly rare (Drury 1974), or simply un-
derrepresented in collections? Two of the
many possible examples illustrate this prob-
lem of identifying endangered bees. Until
1975, Osmia tanneri Sandhouse, a mason bee,
was represented by a single male specimen
collected in 1928 in the Raft River Moun-
tains of Utah by Vasco M. Tanner. F. D.
Parker (1975) rediscovered the species nest-
ing near Wellsville, Utah, and in 1978 Tepe-
dino and Boyce (submitted) found a large
nest in a lawn in Laramie, Wyoming. Fifty
years after the species was discovered we
know little more than that it still exists and it
builds mud nests under rocks.
The genus Dufourea, a ground-nesting
group, provides numerous examples of spe-
cies with restricted distributions. G. E. Bo-
hart of the Logan Bee Laboratory is cur-
rently studying the systematics of this group
and has kindly furnished the following infor-
mation. Over half of the 70 known species
are restricted to California, and many of
these have been recorded from only a single
county. An undescribed species is restricted
to the hills west of San Bruno, an area which
is likely to undergo considerable devel-
opment in the near future. Another undes-
142
Great Basin Naturalist Memoirs
No. 3
cribed species is represented by two speci-
mens collected in Joshua Tree National
Monument. Diifourea macswaini Bohart has
been collected only from the flowers of
Clarkia purpurea in Madera County. Ten to
fifteen other species have restricted, allopat-
ric distributions on the west slope of the
Sierras. With the continuing increase in pop-
ulation and habitat destruction in California,
it is likely that many of these rare species
will disappear.
For a single region, Wyoming shortgrass
prairie, we know that species abundance
curves for bees show the typical insect pat-
tern (Williams 1964): there are a few abun-
dant species and many rare ones (Fig. 1).
Some of this rarity is undoubtedly due to in-
adequate sampling or to the capture of errant
individuals which are abundant at higher ele-
vations 5-10 km away. However, many of
these species may be fugitives (see below),
whose local abundance shows much spatio-
temporal variability.
The Island Effect
With increasing loss of habitat, many plant
and pollinator species will be confined to is-
land preserves of restricted size surrounded
by unsuitable areas. The number of species
supportable will be determined by size of the
preserve and the distance to other preserves,
expressed through immigration and extinc-
tion rates (MacArthur and Wilson 1967).
Such "mainland island preserves" and their
appropriate design have been the subject of
much discussion (Diamond 1975, 1976, Wil-
son and Willis 1975, Simberloff and Abele
1976, Whitcomb et al. 1976, Pickett and
Thompson 1978), but, in general, preserves
should be as large as possible so as to reduce
the probability of extinction of resident spe-
cies. Large size preserves become more im-
portant as suitable surrounding habitat di-
minishes, because immigration rate decreases
with increasing distance from potential
source areas.
The islandlike nature of preserves will also
influence the kinds of plants and pollinators
which can survive. Preliminary studies of the
Galapagos and other islands (Carlquist 1974)
suggest that a depauperate pollinator fauna
restricts successful colonization to auto-
gamous and anemophilous plant species and
to those compatible with the pollinator
fauna. On our size-restricted mainland islands
the flora and fauna is, for the most part, al-
ready present. The questions are: Which por-
tion of the pollinator fauna is most vulner-
able to extinction, which species or species
types are most likely to recolonize, and how
will this affect plant community composi-
tion?
Plant community composition can be al-
tered by differential changes in any of the
numerous selective pressures that operate at
each stage in the life cycle of component
plant species (Harper 1977). Reduction in
seed set, because of changes in the pollinator
fauna, is but one way to alter the abundance
of a given plant species. Nevertheless, polli-
nation is a critical step in the production of
the sporophyte generation. Other factors that
affect the relative success of a plant species,
and ultimately commvmity composition itself;
operate subsequent to pollination. If the
gametophytes are not brought together, no
other factor is important.
The effect of a change in the pollinator
fauna on particular plant species will depend
upon how specialized in pollinator require-
ments a plant is and which species of pollina-
tor(s) has been lost. Specialized species have
long been thought to be less adaptable to
changing conditions and therefore more vul-
nerable to extinction (Rensch 1959). Recent
evidence supports this idea (Drury 1974,
Case 1975, Diamond 1975, Wilson and Willis
1975). A plant which has evolved with a spe-
cific pollinator is doubly disadvantaged be-
cause it may become endangered through di-
rect means, e.g., habitat loss, or by dis-
appearance of its pollinator. Specialized pol-
linators are exposed to similar risks.
Certain specialized pollinator species will
require large tracts of land for their survival.
Traplining species (Janzen 1971), which may
be abundant in the tropics, require large
areas for successful foraging and probably
would not survive on smaller preserves. Plant
species dependent upon traplining pollinators
for outcrossing are also vulnerable.
If floral production by the food plant of a
specialized pollinator is spatiotemporally
variable, then reserves of a size sufficient to
incorporate such variability will be necessary
1979
The Endangered Species: A Symposium
BOULDER RIDGE
143
50-
1975
40-
30-
20-
SPECIES
O
1
Ll
O
L^ ^
1976
u
CD
Z)40-
30-
DIRT FARM
20
10
1 2 3 4 5 6
7 8 9 1
OCTAVE
2 3 4 5 6
Fig. 1. Species abundance distributions for the bee fauna on two rangeland sites in southeastern Wyoming. Each
site-year is treated separately. An octave is equivalent to log2 number of individuals. All curves, except Boulder
Ridge 1975, fit a lognormal distribution.
144
Great Basin Naturalist Memoirs
No. 3
if pollinator and plant are to be preserved.
Little is known about variability in flower
production between years in the same area or
between areas within years. On shortgrass
prairie, spatiotemporal variability in flower
production is substantial (Tepedino and Stan-
ton, in prep.). Other evidence from the liter-
ature tentatively suggests that many other re-
gions exhibit similar variability between
years in floral phenology (Tepedino and
Stanton, in prep.) and that specialized plant
species are no less variable than others (Tepe-
dino and Sauer, unpubl. ms.). If we are to de-
sign natural reserves with necessary min-
imum dynamic area (Pickett and Thompson
1978) to accommodate pollinators, much ad-
ditional data on variability of flower produc-
tion by specialized plant species will be re-
quired. In general, however, smaller reserves
are probably more stressful to specialist
plants and pollinators than to generalized
species.
Regions which harbor high proportions of
specialized pollinators are especially prone to
species loss. The Sonoran Desert, where Neff
et al. (1977) estimated that 33-50 percent of
bee species are specialized, is one such re-
gion. Indeed, much of the bee fauna of a sub-
stantial portion of the western United States,
as well as other arid regions, may be special-
ized (Linsley 1958, Moldenke 1976). If spe-
cializations of euglossine bees are indicative
of bee-plant relationships in the American
tropics (Janzen 1971, Williams and Dodson
1972) then a substantial portion of the tropi-
cal flora and fauna may be jeopardized. Al-
ternatively, the numerous species of stingless,
social bees in the American Tropics, most of
which are probably generalized in flower uti-
lization, may indicate lower diversity of flor-
al reproductive adaptations than we cur-
rently think. Many species of canopy trees,
for example, produce large numbers of small,
nonspecialized flowers (Frankie 1975) that
superficially appear capable of utilizing a va-
riety of insect species. We will need more
data on tropical pollination systems before an
adequate assessment can be made.
Most plant and pollinator taxa are not so
specialized. For example, on Wyoming's
shortgrass prairie most plant species are vis-
ited by many potential bee pollinators and
most bee species utilize several flower spe-
cies (Fig. 2). Moldenke (1975, 1976) reported
similar results for several plant communities
in the western United States. On shortgrass
prairie, flower and pollinator usage also var-
ied widely between years. Using Sorenson's
presence-absence similarity index (Mueller-
Dombois and Ellenberg 1974), we compared
the plants foraged upon for each species of
bee between years and, also, the pollinators
which visited each species of plant in each
year (Fig. 3). Most species of bees and plants
were variable in their resource usage (low
similarity values), especially at the Boulder
Ridge site where floral variation was also
greatest (Tepedino and Stanton, in prep.). Fi-
nally, most bee species also utilized a variety
of flowers during particular foraging trips.
Identification of pollen species from loads
carried by bees showed that over 65 percent
of all individuals had visited two or more
plant species on a given trip and that 46 per-
cent had visited more than three plant spe-
cies.
Generalized bees are less vulnerable to ex-
tinction than specialists for reasons related to
their ability to utilize a variety of flower spe-
cies. First, unlike specialists, generalists will
not become endangered because of the dis-
appearance of a specific host plant. Second,
the probability of a species becoming extinct
due to random events increases with decreas-
ing population size (MacArthur and Wilson
1967). Populations of resident generalists
should be better buffered against wide fluc-
tuations in numbers because of the wider po-
tential resource base. In particular, popu-
lation size of generalist species during
unfavorable periods of bloom should be high-
er than that of specialists and therefore less
prone to extinction. Finally, generalists are
less dependent upon the size of preserves
than specialists, because it is more likely that
surrounding areas will contain plants which
are suitable to them. In effect, the area suit-
able to generalists will almost always be
greater than that for specialists and will ex-
tend beyond the confines of a preserve. Pre-
serves should be designed with the minimum
dynamic area (Pickett and Thompson 1978)
necessary for specialist survival in mind.
One particular group of generalized bees
that is least likely to be affected by habitat
loss and disturbance is the fugitive species
1979
The Endangered Species: A Symposium
145
contingent. Hutchinson (1951) proposed the
term fugitive to describe species that avoid
vying for limited resources with superior
competitors by dispersing to locaHzed patch-
es of resource abundance where competition
is temporarily relaxed. Fugitive bee species
should be especially evident in habitats
where floral resources are spatiotemporally
unpredictable such as shortgrass prairie
(Tepedino and Stanton, in prep.). Indeed, in
these studies on shortgrass prairie, we found
that less than 30 percent of the approx-
00
LJ
LiJ15
CL
OO
cr
UJ
5-
DIRT FARM
a
BOULDER RIDGE
b
5 6-10 11-15 16-20 1-5 6-10 11-15 16-20 >20
NUMBER OF BEE SPECIES VISITING
UJ
0 15-
LJ
Q_
Ul
LJ
LJ
CQlO-
1-2 3-4 5-6 7-8 9-10 =>10 1-2 3-4 5-6 7-8 9-10 11-15
NUMBER OF FLOWER SPECIES VISITED
Fig. 2. a,b— Distribution of shortgrass prairie flower species according to the number of bee species visiting them
over two years on two sites in southeastern Wyoming; c,d— distribuion of bee species by the number of flower spe-
cies utilized.
146
Great Basin Naturalist Memoirs
No. 3
100
80
y 60
u
UJ
Q-
^ 40-
UJ
UJ
CD
5^ 20-
DF BEE SPECIES
a
x=O502 (±.229)
BR BEE SPECIES
b
x = 0.407 (±.219)
BETWEEN YEAR SIMILARITY IN FLOWERS VISITED
DF FLOWER SPECIES
lOOi
80- rr
60
40-
20-
x=0375(±.316)
BR FLOWER SPECIES
d
x"=Q200(±.187)
■75
0- >25- >50- >.75 0- >25- >.50-
.25 .50 .75 .25 .50 75
BETWEEN YEAR SIMILARITY IN BEE VISITATION
Fig. .3. a,b— Distribution of resident bee species by their similarity in flower utilization between consecutive years
on two shortgrass prairie sites; c,d— distribution of flower species by their between-year similarity in bee species
visiting them.
1979
The Endangered Species: A Symposium
147
imately 150 bee species recorded on each of
two sites could meet such relaxed require-
ments for residency as >3 individuals on a
given site in each year. Most species were ei-
tlier present in very low numbers in each
vear or were abundant in one year and ab-
sent in the other. The percentages of total in-
dividuals collected that were nonresidents
varied from 8.6-40.4 percent at the Boulder
Ridge site and from 16.9-26.8 percent at the
Dirt Farm. Although some nonresidents were
undoubtedly incidentals from other habitats,
the large number of species and individuals
in this category suggest the presence of a sub-
stantial fugitive species contingent.
With diminution of native habitat, fugitive
species will comprise an increasingly domi-
nant element of the bee fauna. Unlike spe-
cialized bees and to a lesser extent resident
generalists, fugitives will be uninfluenced by
size of reserves because of their tendency to
disperse and their ability to utilize a variety
of floral resources. Fugitives will have little
difficulty succeeding outside reserves because
of the many patches of floral resources avail-
able along roadsides, in agricultural fields,
backyards, etc. Indeed, these are the kinds of
situations to which fugitives are adapted
(Wilson and Willis 1975, Diamond 1976,
Whitcomb et al. 1976). As resident bee spe-
cies gradually disappear from reserves be-
cause of random extinctions of their relative-
ly small populations, they will be replaced
not by other immigrants of the same species
but by fugitives.
The affect on the flora of losing general-
ized pollinators is difficult to assess. It is
tempting to claim that many generalists are
functionally redundant and therefore expen-
dable; if lost, their pollinatory activities will
be assumed by others. Such a justification for
nonpreservation is potentially insidious be-
cause we have virtually no information on
the relative efficiencies of different pollina-
tors on particular plants or of a single pol-
linator on several plant species (Primack and
Silander 1975). Conversely, several studies
have now shown that plant and pollinator di-
versity are significantly correlated (Heithaus
1974, Moldenke 1975, del Moral and Stand-
ley 1979). A reduction in the diversity of ei-
ther plants or pollinators may lead to a re-
duction in the diversity of the other.
There are reasons for believing that func-
tional redundancy is minimal, and that elimi-
nation of generalized bee species as well as
specialists can lead to differential alterations
in seed set between plant species. First, some
bees may collect nectar and/or pollen from
certain species of flowers without pollinating
them (Grant and Grant 1965, Faegri and van
der Fiji 1971, Percival 1974, Tepedino 1975).
Small species and certain bumble bees are
more likely to fall into this "robber" category
(Faegri and van der Fiji 1971). Their cate-
gorization as potential pollinators of those
plants from which they rob is misleading and
can lull us into a false sense of redundancy.
We must be careful to distinguish between
visitors and pollinators to arrive at intelligent
conservation decisions, and this will necessi-
tate much additional study. Nor is it valid to
conclude that because a generalized pollina-
tor robs the resources of one plant species it
is without value as a pollinator of other spe-
cies. Many bumble bees rob nectar from cer-
tain plants but are important pollinators of
others (Faegri and van der Fiji 1971).
Secondly, although generalized bee species
utilize a broader subset of available floral re-
sources than do specialists, the foraging of
any given species does not include all avail-
able flower species. Neither are the visits of
any particular species proportional to the
abundance of flower species available (Tepe-
dino and Stanton, in prep.), nor are bee spe-
cies equally efficient at pollinating all plant
species visited. The extinction of a single pol-
linator species will reduce visitation rates to
certain plant species in the community to
some unknown degree. Without evidence, it
seems overly optimistic to assume that such a
reduction will be compensated for by re-
maining species. Further, even if visitation
rates by other species do compensate for the
lost pollinator, there is no basis for assuming
that the efficiency of such visits is equivalent
to that of the species which have dis-
appeared. For example, in a study of four
solitary bee species visiting alfalfa, Batra
(1976) found that, although all gathered nec-
tar and pollen and accomplished pollination,
they did so with varying degrees of pro-
ficiency. Two species spent more time than
others foraging on hidden flowers, one visited
many more flowers which had already been
148
Great Basin Naturalist Memoirs
No. 3
pollinated, etc. The species varied during
morning foraging from 0.8 to 3.68 pollina-
tions per minute. The effect of removal of
one of these pollinators upon seed set would
depend upon which species was removed. If
we extend the results of this simple green-
house study to plant communities in the field,
we gain an impression of the unknown com-
plexities which we are tampering with.
It seems clear that plant species that de-
pend upon particular bees for their reproduc-
tion will experience severe selective pressures
to evolve autogamy (Levin 1972) or wind
pollination or to realign their floral morpho-
logies to take advantage of remaining pol-
linator species. For many specialized plants
such adaptations will be impossible (Levin
1971, Baker and Hurd 1968) and their extinc-
tion is likely. While it is not uncommon to
develop facultative autogamy from obligate
outcrossing (Baker 1959), selling may be least
advantageous in environments with low pre-
dictability such as shortgrass prairie (Solbrig
1976). In such regions plant species that solve
pollination problems with obligate autogamy
may become extinct more gradually.
As specialized pollinators are replaced by
fugitives, more generalized plant species may
become endangered because fugitives are rel-
atively inconstant foragers. Several theo-
retical studies provide similar results for situ-
ations in which plant species compete for
pollinator visits: if pollinator constancy is
proportionate to floral abundance, minority
species will receive fewer pollinating visits
than more abundant species and will eventu-
ally disappear (Levin and Anderson 1970,
Straw 1972, Waser 1978). Even worse, if pol-
linators show disproportionate preference for
more abundant species, then less abundant
species will approach extinction more rapid-
ly-
There is little doubt that in North America
we will lose many bee species and other pol-
linators as well, particularly from the western
states. As a result of these extinctions, we will
probably see some gradual transition in the
composition of our flora. Floral change will
be most obvious and far-reaching in desert,
chaparral, and alpine ecosystems, where the
percentage of insect-pollinated plants is high
(Moldenke 1976). In forests and grasslands,
where the dominant plant species are wind
pollinated, changes will be more subtle and
less easy to predict.
Obviously, the key to slowing the rate of
pollinator and plant extinction is habitat
preservation. We need to set aside as much
land as we can possibly afford in the form of
greenbelts, parks, and reserves of various
sizes. In addition, we should encourage the
use of local plant species as ornamentals in
backyards and gardens instead of the sterile
creations of seed companies. Local plant spe-
cies are frequently as esthetically pleasing
and, because they are adapted to the region,
require less care and expense in the form of
fertilizers, water, etc. It is also quite simple
to provide nesting material for some solitary
bees in the form of pine wood blocks or
scraps with holes drilled in them (Krombein
1967). These can be set out on posts in back-
yards as are bird houses and feeders. The spe-
cies which will utilize these trap-nests are not
at all agressive and will sting only when han-
dled. In short, every little bit will help and,
unfortunately, we need all the help we can
get.
Acknowledgments
F. D. Parker and P. F. Torchio provided
helpful suggestions on the manuscript. I
thank G. E. Bohart for unpublished informa-
tion on the genus Dufourea and B. B. Becker
for the figures.
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ENDANGERED SPECIES: COSTS AND BENEFITS'
Edwin P. Pister-
.\bstract.— Biologists are often placed in the difficult position of defending a threatened habitat or animal with
vagiie reasoning and faulty logic, simply because they have no better rationale at their immediate disposal. This
places them at a distinct disadvantage and literally at the mercy of resource exploiters and their easily assignable
dollar values. Although the initial dollar cost of delaying or precluding "developing" may be significant, the long-
term benefits of saving the biological entities which might otherwise be destroyed are likewise great and are measur-
able in concrete terms which society is only now beginning to appreciate. Case histories are presented, a more pro-
found rationale is explained, and the environmentalist is challenged to make his case sufficiently effective to reverse
the current exploitive trends which threaten so many of earth's life forms.
The land and water developers, mineral
extractors, and other resource users which
burgeoned nationwide (especially in the
West) following World War II placed agency
resource managers in a new and uncomfort-
able position. Whereas there once had been
sufficient land and water for everyone, in-
cluding our plant and animal species, we sud-
denly found ourselves entering into what
seemed (on the surface, anyway) an "us or
them" situation. Backed into a corner, biolo-
gists and administrators found themselves
searching frantically for values with which to
defend their trust against the hard dollar fig-
ures of the exploiter.
Nowhere has this concept been more ap-
parent than in our efforts to preserve threat-
ened and endangered species. When pitted
against a potential development project in-
volving the expenditure of millions of dollars,
the environmentalist has been forced to bol-
.ster his innate sense of doing what he knows
is right with whatever biological rationale
might enter his mind. Often his reasoning
proves biologically unsound, reducing his
changes of success and injuring his profes-
sional credibility.
A ray of hope has been noted recently
through the presentation of a new rationale,
one which bolsters valid existing arguments
with profound .spiritual values. This paper
presents a brief history of recent preservation
efforts, summarizes the new rationale, and of-
fers the hope that newly defined goals, al-
though lofty, are by no means unattainable.
Acknowledgments
It would be ungrateful of me to prepare a
paper on endangered species, especially one
involving fishes, without acknowledging the
enormous efforts of Professor Carl Hubbs of
Scripps Institution of Oceanography and Dr.
Robert Rush Miller of the University of
Michigan. Many others, agency biologist and
academician alike, have made great contribu-
tions to the cause, especially in recent years.
Special thanks are due Dr. David W. Ehren-
feld of Rutgers University, Dr. Hugh W. Nib-
ley of Brigham Young University, and Mr.
Jimmie Durham of the International Indian
Treaty Organization, whose works I have re-
ferred to and quoted extensively in the prep-
aration of this paper.
Background
My first real involvement with endangered
species began in July 1964 during a field trip
with Bob Miller and Carl Hubbs in Califor-
nia's Owens Valley. Bob suspected that the
Owens pupfish {Cijprinodon radiosus) was ex-
tinct when he described it (Miller 1948), but
he felt it worthwhile to make one final effort
'Initially presented at a symposium. Fishery Benefits to Society and Industry, as part of the joint annual conference of the Western Association of Fish
and Wildlife Agencies and Western Division American Fisheries Society, San Diego. California, 17-20 July 1978.
■California Department of Fish and Game and Executive Secretary, Desert Fishes Council, 407 West Line Street, Bishop, California 93514.
151
152
Great Basin Naturalist Memoirs
No. 3
to locate a remnant population. My interest
in endangered and nongame fishes was min-
imal at that time; yet, when exultant voices
resounded through the marsh, a strange feel-
ing came over me. My conversion to the
cause of nongame species was instantaneous
and dramatic. Fish Slough on the floor of
Owens Valley was my Road to Damascus
(Miller and Pister 1971), and I have been an
active crusader for the past 14 years, almost
to the day.
Sympathy for the cause was rare within
my agency, and time and funds were vir-
tually imavailable for anything but the man-
agement of game species. What work we
were able to accomplish was generally done
on our own time and expense.
Yet somehow the movement grew. Support
was excellent from the academic community;
and the Fish and Wildlife Service, which
found itself less encumbered by politics and
tradition than those of us in state agencies,
also offered good assistance, often to cries of
anguisH concerning the "Federal Octopus"
from within Great Basin state directorships.
Sadly, this point of contention still precludes
optimum interagency management program-
ming, and we still eagerly await the day
when welfare of the resource will overcome
agency jurisdiction as the primary point of
concern.
The initial meeting of the Desert Fishes
Council, formed in desperation in 1969 to
stave off the almost certain extinction of sev-
eral fishes within the Death Valley drainage
system, drew 44 individuals, primarily with
federal or academic affiliations (Pister 1974).
The 1978 Council membership approaches
300 and is growing rapidly as public recogni-
tion of the need for desert ecosystem pro-
tection increases.
My involvement in endangered species
work (and colleagues often state the same
motivation) stems from a desire to leave
something significant as a mark of my having
been here. Somehow, in my advancing ca-
reer, the idea of providing a bunch of game-
fish for people to catch simply was not suffi-
ciently fulfilling. It became apparent to me
that if man were ever to exercise his domin-
ion (a term which until recently was disturb-
ingly vague) in an acceptable way, he was
going to have to turn a new leaf and face a
new set of problems.
Discussion
At a recent symposium on endangered spe-
cies held at Yale University, Dr. Lee Talbot,
vice-president of the International Union for
the Conservation of Nature, said that even
obscure endangered species can serve as in-
dicators of large environmental problems
that may have major adverse effects on
people who could not care less about the ani-
mal in question. In this context, let us consid-
er the following examples:
Devils Hole
East of Death Valley, Nevada, is a lime-
stone cavern with a tiny pool containing the
entire world population of the Devils Hole
pupfish (Cyprinodon diaholis). Extensive and
indiscriminate agricultural development and
irrigation in the late 1960s was rapidly
lowering the aquifer system supplying the
pool, and it was apparent that, unless some-
one did something to stop it, a full species
(the most highly evolved of the Death Valley
cyprinodonts) would soon become extinct.
This actually was the cause celehre which
motivated us to form the Desert Fishes Coun-
cil. We fought long and hard in behalf of De-
vils Hole and its inhabitants and found, in the
process, that the State of Nevada, with the
exception of its Department of Fish and
Game, was often uncooperative and even an-
tagonistic when asked for assistance in stop-
ping the deadly water table drawdown. This
seemed particularly true of the state engi-
neer's office. Nevada is very development
oriented and, despite a rather paradoxical
state endangered species law, generally
viewed as highly undesirable a tiny fish of no
economic value which seriously threatened a
multimillion dollar ranching operation.
Federal law seemed to offer our only salva-
tion in this matter, so in July 1972 the People
of the United States, through the Department
of Justice, went to court against the land de-
veloper and the State of Nevada as codefen-
dants. Interestingly, the case was not argued
on the basis of the Endangered Species Act,
but on a point of water law.
1979
The Endangered Species: A Symposiui
153
Initial judgment was favorable, and after a
siege in the appellate court the case was
heard (amidst the strong desire of western
congressmen to reverse the earlier decisions)
by the U.S. Supreme Court. In June 1976 the
court ruled unanimously in favor of the
People of the United States (and the fish),
and we began to regain some confidence in
"the system." It was encouraging to know
that the Equal Justice under Law inscription
over the entrance to the Supreme Court
building in Washington, D.C., applies to fish,
too.
Probing deeper into the matter, we found
that the Ash Meadows ranching operation
was only a symptom of a much greater envi-
ronmental threat. A report by a consultant to
the Nevada State Engineer (Nevada State
Engineer's Office 1971) to locate and eval-
uate future water sources for Las Vegas ear-
marked underground supplies around Devils
Hole to provide 3 million acre feet over a 30-
year period. At this time, the report in-
dicated, it would no longer be feasible to nm
the pumps, and the deteriorating quality of
what remained would make it unsuitable
anyway. However, that 3 million acre feet
would allow a sufficient increase in popu-
lation to facilitate acquisition of water from
more permanent sources farther away (such
as the Columbia or Snake rivers).
In this case the Devils Hole pupfish proved
to be an indicator organism which led, even-
tually, to a discovery of the underlying poli-
tics of the entire matter. Would it be to
man's long-term benefit to destroy a spring
ecosystem unique in the United States and
equaled only in one location in Mexico sim-
ply to provide short-term water to a city
which must very obviously someday curtail
its growth? At least we now have the chance
to take a harder look.
Tellico Dam and the Snail Darter
In a related situation, news media through-
out the United States have recently been dis-
cussing a situation on the Little Tennessee
River wherein the tiny snail darter {Percino
tanasi) has essentially stopped completion of
the $116 million Tellico Dam following a 6-3
Supreme Court decision affirming the provi-
sions of the Endangered Species Act of 1973.
Reaction by the media has been mixed, with
some lauding the decision and others con-
demning an action that, in their estimation,
would waste $116 million simply to save a
fish of no known economic value. In the
wake of this. Congress (with many members
in an election year asking themselves "What
if Tellico were in my district or state?") is
now debating whether or not the Endangered
Species Act should even be renewed and, if
so, what amendments should be made to "al-
low greater flexibility."
Again, a look behind the scenes is reveal-
ing. First off, TVA (the sponsoring agency)
failed to discuss the snail darter problem with
the Fish and Wildlife Service until the proj-
ect was nearly finished, and greatly accelera-
ted the construction schedule to create a
stronger case for completing the dam. Sec-
ondly, a General Accounting Office study
(U.S. General Accounting Office 1977) imple-
mented by Congress revealed a cost-benefit
analysis so faulty that even after the expendi-
ture of over $100 million taxpayers would be
money ahead if the dam were torn down. In
fact, this alternative was offered by TVA
Chairman S. David Freeman before a House
subcommittee following the Supreme Court
decision and release of the General Account-
ing Office report. Lastly, considering the
widespread pressure to terminate or weaken
the Endangered Species Act, it is significant
to note that in all except one (Tellico) of
more than 4,500 consultations between de-
velopers and the Fish and Wildlife Service,
both the project and the species in question
were deemed able to coexist.
Additional Benefits
In addition to the above-listed benefits of
revealing the political issues underlying vari-
ous development proposals, concern over en-
dangered species has resulted in beneficial
philosophical shifts within many state fish
and wildlife management agencies. Although
the primary orientation of such agencies re-
mains one of providing a harvestable product
for hunters and anglers, changes are being
noted through the implementation of non-
game, endangered species, environmental,
and land acquisition programs as the future
154
Great Basin Naturalist Memoirs
No. 3
demands of society become increasingly ap-
parent (Pister 1976).
Costs
What are the direct agency costs of rare
and endangered species programs? Small, by
most standards. In 1978, according to Fish
and Wildlife Service figures, the cost of ad-
ministering the Endangered Species Act, in-
cluding aid to the states, was $16.2 million. A
figure of $19.4 million is estimated for 1979.
It is virtually impossible to accurately assess
the dollar costs of delaying a development
project during the discussion period with the
Fish and Wildlife Service.
A New Rationale
Why do the people of the United States
find themselves in the current dilemma? Per-
haps Nibley (1978:85-86) says it best: "We
have taught our children by precept and ex-
ample that every living thing exists to be
converted into cash, and that whatever
would not yield a return should be quickly
exterminated to make way for creatures that
do." I cannot think of a better way to put it,
and I am reminded of Paul's admonition to
Timothy nearly 2,000 years ago: "For the
love of money is the root of all evil. . . ." (I
Tim. 6:10).
In view of the increasing concern of vir-
tually all segments of society over environ-
mental matters generally, and noting this
same concern within academic circles, it ap-
pears to me highly appropriate that two of
the most outstanding essays involving endan-
gered species to emerge during the past dec-
ade should be written by eminent scholars
representing two very different disciplines.
David W. Ehrenfeld, a Harvard M.D. with a
Ph.D. in zoology and biochemistry from the
University of Florida, is currently professor
of biology at Cook College, Rutgers Univer-
sity. Hugh Nibley graduated in history from
UCLA and received his Ph.D. from the Uni-
versity of California at Berkeley. Adept in 14
languages, he taught history and languages at
the Claremont Colleges in California before
moving to Utah. He is now professor of his-
tory and religion at Brigham Young Univer-
sity.
Yet, although their academic disciplines
may differ, their philosophies blend marve-
lously well, complement one another, and
lead to a logical and acceptable rationale for
the preservation of all life forms.
Ehrenfeld (1976), in a masterful essay en-
titled "The Conservation of Non-Resources,"
does the biologist a great favor by critically
analyzing the most popular (and frequently
contrived) reasons advanced in defense of a
favorite species or program. He defines a re-
source as a commodity that has an appre-
ciable money value to man and then lists sev-
eral that do not. These he considers to be
non-resources, without conjectural or demon-
strated resource value to man. He utilizes the
Houston toad (Bufo houstonensis) to exem-
plify this concept and throws fear into the
hearts of many zealots when he states quite
accurately that certain species may even ex-
hibit a negative value. Ehrenfeld warns
against the dangers of prioritizing, or rank-
ing, species or natural areas in a preservation
program because of our categorical lack of
knowledge about them, be it now or 100
years from now. He feels, further, that formal
ranking sets natural area against natural area
(and species against species) in an unaccept-
able and totally unnecessary way, and em-
phasizes that the need to conserve a particu-
lar community or species must be judged
independently of the need to conserve any-
thing else (Ehrenfeld 1976:653).
He then goes on (p.654) to state that only
one account exists in Western culture of a
conservation effort greater than that now
taking place, where not a single species was
excluded on the basis of low priority, and by
all accounts not a single species was lost
(Genesis 7:8-9):
Of clean beasts, and of beasts that are not clean, and
of fowls, and of everything that creepeth upon the earth,
There went in two and two unto Noah into the ark,
the male and the female, as God has commanded Noah.
It is encouraging to note that even (or per-
haps especially) the more sophisticated
writers seem to be rejecting the classical,
anthropocentric economic arguments for spe-
cies preservation in favor of a religious con-
cept presented by Elton (1958) 20 years ago
and hirther developed by Ehrenfeld
(1976:654-655), who states:
1979
The Endangered Species: A Symposium
155
The non-economic value of coniniunities and species
is the simplest of all to state: they should be conserved
because they exist and have existed for a long time.
Long-standing existence in nature is deemed to carry
with it the unimpeachable right to continued existence.
Existence is the onlv criterion of value, and diminution
of the number of existing things is the best measure of
decrease of value. This is, as mentioned, an ancient way
of evaluating "conservability" and by rights ought to be
named the "Noah Principle" after the person who was
one of the first to put it into practice.
In recent hearings on the Endangered Spe-
cies Act held by the House Merchant Marine
and Fisheries Committee, Jinimie Durham,
director of the International Indian Treaty
Organization, posed a very logical and per-
tinent question: Who has the right to destroy
a species? Because of Durham's eloquence,
any attempt to paraphrase his statements
would markedly reduce the feeling which his
words convey. The following material has
been extracted from his published address
(Durham 1978):
In Ani Yunwiijah, the language of my people, there is
a word for land: Eloheh. This same word also means his-
tory, culture, and religion.- This is because we Cherokees
cannot separate our place on the earth from our lives on
it, nor from our vision and our meaning as a people.
From childhood we are taught that the animals and
even the trees and plants that we share a place with are
our brothers and sisters.
So when we speak of land, we are not speaking of
property, territory or even a piece of ground upon
which our houses sit and our crops are grown. We are
speaking of something truly sacred.
There is no Cherokee alive who does not remember
that Trail of Tears, as we call our march into exile in
Oklahoma. There is none among us who does not re-
member and revere that sacred land, Echota.
Today, the Tennessee Valley Authority would like to
flood the sacred valley that held our two principal cities,
Echota and Tenasi, after which the state is named. The
Tellico project would have destroyed an area of great re-
ligious importance, many settlement sites, cemeteries,
rich farmlands, forests and the river itself. This is an un-
needed dam which can, at the whimsy of TVA, wipe out
thousands of years of history of a great and currently op-
pressed people. To do so would be an insult not only to
the Cherokee, but also to all the people in the United
States and to humanity. Yes, I am proud enough to state
that the history and vision of my people are important
to humanity.
The flooding of our valley has been stopped temporar-
ily because of a little fish that lives there and nowhere
else. I have seen Atty. Gen. Griffin Bell, the New York
Times and a national television network make fim of this
little fish and I would like to ask why it is considered so
humorously insignificant. Because it is little, or because
it is a fish?
It is this incredible arrogance towards other life that
has caused such destruction in this countrv. Who is Grif-
fin Bell or the U.S. government to play God and judge
the life or death of an entire species of fellow beings
which was put here by the same power that put us here?
Who has the right to destroy a species of life, and what
can assuming that right mean?
Let me be emotional: To me, that fish is not just an
abstract "endangered species" although it is that. It is a
Cherokee fish and I am its brother. Somehow, it has
acted to save my holy land, so I have a strong gratitude
for that fish.
The Cherokee people in Tennessee, Oklahoma, the
Carolinas, Georgia and wherever we might be are of one
voice and of one mind that this dam, this degradation,
must be stopped. We want our universe, our Echota
with all of its fish and all of its life to continue. We are
sine that this cannot be against the interests and wishes
of the American people.
Definitions
Although subdue and dominion as used in
Genesis carry a religious connotation, vir-
tually all environmentally oriented dis-
cussions in which these words arise seem to
end with everyone defining them to suit his
own selfish purposes.
We have long been in need of a clear and
learned treatise on this subject, and the entire
cause of species preservation is fortunate in-
deed to have someone of Hugh Nibley's stat-
ure and capability to provide one for us (Nib-
ley 1978:85-99). His analysis of man's
dominion borders on pure genius, and he log-
ically asks in his preface (p.86): "If God were
to despise all things beneath him, as we do,
where would that leave us?" He then pro-
ceeds typically to use a wealth of scripture,
classical literature, and other references to
develop his theme that "Man's dominion is a
call to service, not a license to exterminate"
(p.96), and provides an example from a pio-
neer leader: "while 'subduing the earth' we
must be about 'multiplying those organisms
of plants and animals God has designed shall
dwell upon it,' namely 'all forms of life,' each
to multiply in its sphere and element and
have joy therein." (p.87). This was indeed an
inspired statement from the leader of a group
of pioneers seeking to tame a desert wilder-
ness. Nibley suggests an in-depth analysis of
the derivation of "dominion," which clearly
turns out to be the responsibility of the mas-
ter for the comfort and well-being of his de-
pendents and guests, "not a predator, a ma-
nipulator or an exploiter of other creatures,
156
Great Basin Naturalist Memoirs
No. 3
but one who cooperates with nature as a dih-
gent husbandman" (p.88).
Nibley continues: "The teaching of Israel
laid the heaviest emphasis on responsibility.
Since man is quite capable of exercising the
awesome powers that have been entrusted to
him as the very image of God, he must needs
be an example to all, and if he fails in his
trust, he can only bring upon himself the con-
demnation of God and the contempt of all
creatures." (pp.89-90).
Nibley 's explanation of man's hostility is as
logical and obvious as it is painful: "The ani-
mal, vegetable, and mineral kingdoms abide
the law of their Creator; the whole earth and
things pertaining to it, except man, abide the
law of their creation, while 'man, who is the
offspring of the Gods, will not become sub-
ject to the most sensible and self-exalting
principles.' (Journal of Discourses, 9:246).
With all things going in one direction, men,
stubbornly going in the opposite direction,
naturally find themselves in the position of
one going the wrong way on the freeway
during rush hour; the struggle to live be-
comes a fight against nature. Having made
himself allergic to almost everything by the
Fall, man is given the choice of changing his
nature so that the animal and vegetable crea-
tion will cease to afflict and torment him, or
else of waging a truceless war of extermina-
tion against all that annoys him until he ren-
ders the earth completely uninhabitable."
(pp.94-95).
Summary
The obvious benefits of endangered species
programs may therefore be summarized as
follows:
1. Endangered species generally serve as
indicators of larger environmental prob-
lems and, when detected, allow analysis
and correction of more involved issues
during the pursuit of a preservation
program.
2. The "Era of Endangered Species" has
initiated a process of maturation within
state fish and wildlife agencies as they
begin to consider all species in their
program planning, not simply those
with an obvious economic value.
3. By preventing the extinction of fish and
wildlife species (and all life forms), we
automatically preserve any anthropo-
centric values which they may possess,
but which research may not yet have
discovered.
4. Perhaps the most important reason for
preserving endangered species is the re-
alization of the opportunity granted to
man— the only species endowed with
the capability of truly caring for his fel-
low creatures— to exercise righteously
the dominion granted him by his Crea-
tor. Doing so will do much to preserve
man's self-respect. The manifestations
of this concept can be enormous, in-
cluding peaceful coexistence with na-
ture, other nations, and himself.
Conclusion
Considering our rather dismal record to
date, including threatened changes in the En-
dangered Species Act resulting from the Tell-
ico Dam-snail darter conflict, the cynic
would consider it quite improbable that man
would ever categorically accept a religious
(or morally based) reason for preserving other
life forms. At this point I must assume the
role of the optimist and state that a widely
accepted nonresource rationale is not only
desirable, it is absolutely mandatory if we are
ever to gain the necessary political strength
to assure adequate recognition of the biota in
a proposed development project. It seems un-
likely in the foreseeable future that, in terms
of dollars, we will ever be able to place a
higher value on the Devils Hole pupfish than
on a section of resort condominiums in Las
Vegas, or prove that the snail darter swim-
ming above Tellico Dam has an economic
worth in excess of the electricity produced by
the water in which it lives.
Ehrenfeld (1976) states quite correctly that
if nonresource arguments are ever to carry
their deserved weight, cultural attitudes will
have to be changed. Tliis is a big order, but
we have no alternative but to try. Henry
Ford used to remind his plant managers:
"You can say it can be done, or you can say it
can't be done and be correct either way."
An analysis of Section 2 (Findings, Pur-
poses, and Policy) of the Endangered Species
1979
The Endangered Species: A Symposium
157
Act of 1973 indicates that Congress appar-
ently felt it was worth a try to implement
such a cultural change, inasmuch as the states
(often the hardest to sell in such matters) and
other interested parties are encouraged to de-
velop and maintain conservation programs
which meet national and international stan-
dards as a key to better safeguarding, for the
benefit of all citizens, the nation's heritage in
fish and wildlife. Further, the purposes of the
act include providing a means whereby the
ecosystems upon which endangered and
threatened species depend may be conserved.
Lastly, the policy of Congress is stated that
all federal departments and agencies shall
utilize their authorities in furtherance of the
act. Although the act lists the physical means
of achieving its purposes, it fails to address
the matter of enlisting and sustaining philo-
sophical support. Inasmuch as the long-term
effectiveness of any legislation is dependent
upon its acceptance by the people, it is im-
plicit that the major responsibility for assur-
ing this falls upon those of us who feel
strongly about such things.
Reflection
Not long ago I arose early and went for a
walk near my Bishop home. I glanced west-
ward and watched the moon set just as the
first rays of the rising sun began to tint the
great peaks of the Sierra Nevada crest. The
effect was spectacularly beautifid and, to me,
illustrated the concept of "the beginning and
the end. " The beginning was represented by
an unprecedented degree of enlightenment
within the American public and in our own
philosophies, and a renewed ability as
agencies and individuals to work together to-
ward the management and preservation of all
of the nation's (and world's) life forms; the
end by a lessening and ultimate cessation of
the anthropocentric attitudes within the pub-
lic and ourselves which have in so many in-
stances "come home to roost" and caused our
current dilemma.
The sun continued to rise and the red turn
to gold as my thoughts went back to the
early days of our desert fish programs. How
utterly hopeless everything seemed then! I
uttered a silent prayer that the insight, hard
work, and example of the earliest workers in
this field might inspire us to better serve the
multitudes who will come after, and that we
might provide them with a legacy reflecting
not only our scientific competence, but also
our practicality and philosophical maturity;
and that this in turn would constitute a cross-
roads in American thought concerning man's
dominion over the earth, and recognizing the
absolute truth that the glory of God is in-
telligence, I ended my prayer with a plea
that we might utilize our collective in-
telligence to glorify Him by exercising a
truly righteous dominion equally over His en-
tire creation.
It seems fitting to express here the
thoughts of the late anthropologist and hu-
manist Loren Eiseley (1962, preface): "I be-
lieve in Christ in every man who dies to con-
tribute to a life beyond his life." He
continues: "I have been accused of woolly-
mindedness for entertaining even hope for
man. I can only respond that in the dim
morning shadows of humanity, the in-
articulate creature who first hesitantly
formed the words for pity and love must
have received similar guffaws around a fire.
Yet some men listened, for the words sur-
vive."
And the Devils Hole pupfish and snail
darter survive, too. Twenty years ago they
wouldn't have had a chance.
Literature Cited
Durham, J. 1978. Who has the right to destroy a spe-
cies? Los Angeles Times, 2 July 1978. Adapted
from a statement delivered before House Mer-
chant Marine and Fisheries Committee.
Ehrenfeld, D. W. 1976. The conservation of non-re-
sources. American Scientist 64(4):648-656.
Eiseley, L. C. 1962. The immense journey. Time, Inc.
Book Division, reprinted by Random House. 152
pp. Time, Inc., New York.
Elton, C. S. 1958. The ecology of invasions by animals
and plants: 14.3-45. London: Methuen.
Journal of Discourses. Deseret Book Company, Salt
Lake City, Utah. 27 volumes, 8th reprint, 1974.
Miller, R. R. 1948. The cyprinodont fishes of the Death
Valley systems of eastern California and south-
western Nevada. Misc. Publ. Mus. Zool. Univ.
Mich. 68: 1-15,5.
Miller, R. R., and E. P. Pister. 1971. Management of
the Owens pupfish, Cyprinodon radiostis, in
Mono County, California. Trans. Amer. Fish.
Soc. 100 (,3): 502-509.
158
Great Basin Naturalist Memoirs
No. 3
Nevada State Engineer's Office. 1971. Water supply
for the future in southern Nevada. Special plan-
ning report. Prep, by Montgomery Engineers of
Nevada. 89 pp. plus references, bibliography,
map and photographs. Nevada State Engineer's
Office, Carson City.
NiBLEY, H. W. 1978. On subduing the earth, pp. 85-99.
In: Nibley on the timely and the timeless. Vol. 1,
Religious Studies Monograph Series, Religious
Studies Center, Brigham Young University. Pub-
lishers Press, Salt Lake City, Utah. 32.3 pp.
PisTER, E. P. 1974. Desert fishes and their habitats.
Trans. Amer. Fish. Soc. 103(3):531-540.
1976. A rationale for the management of non-
game fish and wildlife. Fisheries (Bull. Amer.
Fish. Soc.) 1(1);11-14.
United States General Accounting Office. 1977.
The Tennessee Valley Authority's Tellico Dam
project— costs, alternatives, and benefits. Report
to the Congress by the Comptroller General of
the United States, October 14, 1977.
Warren, C. 1978. Our economics is too small. Unpub-
lished keynote address delivered at joint confer-
ence of Western Association of Fish and Wildlife
.Agencies and Western Division ,\merican Fish-
eries Society, San Diego, California. 18 July 1978.
ENDANGERED SPECIES ON FEDERAL LANDS
PANEL: PART I,
INTRODUCTION
John L. Spinks'
Since I've already spoken once during the
symposium, I only have two brief points to
make for my part of the panel presentation.
One is in terms of public land. The Fish
and Wildlife Service has about 35 million
acres in the National Wildlife Refuge System.
The management of those resources are sub-
ject to the same Section 7 scrutiny as any
other federal agency action. As a matter of
fact, by policy from the director of the Fish
and Wildlife Service, it is our responsibility
to make certain that we live up to the high-
est expectations in compliance with Section
7. If there is a finding of either adverse modi-
fication of critical habitat or a jeopardy find-
ing, that activity will not be done by the Fish
and Wildlife Service— and that is in writing
from the director.
The second point I would make is that,
though the Fish and Wildlife Service has a
lead agency role, as does the National Marine
Fisheries Service, in administering the En-
dangered Species Act of 1973 as amended, I
hope all of you here can immediately grasp
that the job of protecting endangered and
threatened species and recovering these spe-
cies is completely beyond the scope of any
one agency. Were it not for the real dedica-
tion and assistance that the service gets from
folks like these up here and their agencies,
not to mention all the 50 state agencies and
the very concerned and dedicated private in-
dividuals, we would never get to first base.
As a matter of fact, on behalf of the service, I
think all we can say is we appreciate the as-
sistance we've gotten over the years— it has
been continuous and is still forthcoming— and
the interest that generates a symposium like
this. We certainly appreciate the attendance
of all those here.
PANEL: PART II,
FOREST SERVICE PHILOSOPHY OF ENDANGERED SPECIES MANAGEMENT
Jerry P. Mcllwain'
We have heard some excellent talks on en-
dangered species philosophy here at this ses-
sion, treating strategies, genetics, ecology,
and some new techniques and concepts that
are very interesting to me. Within the limita-
tions that are placed on a federal agency, the
Forest Service has been dealing with many of
these philosophies and strategies for a long
time. We have been trying to get them down
to the ground level and convert these things
that we have all been talking about for the
last day and a half into on-the-ground man-
agement, and that is basically what I am go-
ing to talk to you about today.
I will talk about the Forest Service philos-
ophy of endangered species management and
how this policy is being translated into poli-
cies and procedures to get the job done,
about the overall program to accomplish our
endangered species job, land management on
the national forest system, and how the re-
search and state and private forestry arms of
the Forest Service are affected by the Endan-
gered Species Act.
'Chief, Office of Endangered Species, U.S. Fish and Wildhfe Service, Washington, DC. 20240.
'Endangered Species Speciahst, USDA, P.O. Box 2417, Washington, DC. 20013.
159
160
Great Basin Naturalist Memoirs
No. 3
The Forest Service has been in the endan-
gered species game for a long time. We set
up the Sespe Condor Sanctuary on the Los
Padras National Forest in 1946 and had been
studying this bird for a considerable number
of years before that.
Programs to protect and manage bald
eagles, ospreys, Kirtland's warblers, and sev-
eral others were implemented on national
forest system lands long before the Endan-
gered Species Act was passed. Passage of the
act in 1973 did give our program consid-
erable impetus and made endangered species
management an organic part of our agency
responsibilities.
The evaluation of policy and procedures
for the management of endangered species is
very dynamic at the moment because things
are changing so rapidly. Knowledge of the bi-
ology of listed species is being acquired rap-
idly, Congress has recently amended the law,
and Fish and Wildlife Service regulations are
continually evolving. We have been trying to
get a new Forest Service Manual chapter out
now for almost three years. It was just about
ready to go before the endangered species
amendments of this year were passed which
did away with some of our policies and pro-
cedures, so we are back to the drawing
board.
The basic Forest Service philosophy of en-
dangered species management is to meet
both the letter and the spirit of the law by
achieving the recovery of listed species on
national forest lands, not jeopardizing listed
species in our other programs, and assuring
that Forest Service management does not
contribute to a sensitive species qualifying
for listing.
The Forest Service moved out rather rap-
idly in establishing a positive program after
the 1973 act was passed. We feel that our
programs are going a step further than the
requirements of the law in many cases and
are establishing a comprehensive endangered
species program.
Our program considers not only the feder-
ally listed species, but also state-listed spe-
cies, plus a third category we are calling sen-
sitive species. Sensitive species are those
which are proposed to be federally listed or
species that are recognized by the Regional
Forester to need special management in order
to prevent the need for their placement on
federal or state lists. All plants that have been
officially proposed to be listed are considered
sensitive and managed as if they were already
listed.
Some interpretations of the law are that
the legal requirements exist only as long as a
species is listed. If recovery were achieved
for that species and it were removed from
the list, there would be no more legal pro-
tection for that species. The species could
then decline to the point that it was relisted
and the cycle would begin again. Our pro-
grams are aimed at achieving recovery of a
listed species and continuing that status in
perpetuity.
Endangered species program changes have
outstripped the flexibility of our budgeting
system. The Forest Service budget is gener-
ated at the ground level and aggragated up-
ward. It is also formulated two years ahead of
time. The Endangered Species Act Section 7
regulations were just finalized this past Janu-
ary and they impose a considerable number
of requirements on the Forest Service and
other federal agencies. Of course, our budget
was already formulated and was not respon-
sive to the increased work load brought
about by the new regulations. Considerable
budget adjustments made in the Washington
office were necessary. To avoid this hap-
pening in the future, it was necessary to pre-
pare two national programs, one for plants
and one for animals.
The general thrust of these programs is in
three phases: inventory, interim manage-
ment, and recovery management. These
three phases relate to each individual species
of concern. We are in phase 1 for a certain
group of species, phase 2 for another group,
and phase 3 for others.
Basically the first phase is analysis of the
situation: identification of research needed,
really finding out where we are on a species,
and what we need to do. Then we move into
the second phase, an interim management
phase. This is the actual conducting of
needed research relative to habitat require-
ments, establishing management programs,
and protecting the species while we are
doing this. The third and final phase is recov-
ery management. This final phase is initiated
after recovery plans or other specific Forest
1979
The Endangered Species: A Symposium
161
Service plans have been prepared to protect
the species. The species and their habitats are
managed to achieve recovery and prevent re-
currence of endangernient.
Because this is a panel on public land man-
agement, we will now turn to some of the
specific programs on the national forest sys-
tem lands.
Of the 236 domestic species currently on
the federal list, there are about 70 species
that occur on national forest system land. Of
these 70, there are quite a few that occur
only on Forest Service lands or Forest Ser-
vice lands play an essential part of the total
conservation effort for that species. The 1973
act considerably changed the way we do
things in the national forests. The act re-
quired us to evaluate all the Forest Service
projects, decide whether or not they may af-
fect a species, and, if so, enter into the formal
consultation process with the Fish and Wild-
life Service. This has been a considerable
work load and will certainly grow larger in
the future. We have had well over a hundred
formal consultations since the regulations be-
came effective in January of this year. Some
of them have been very complex. Our field
people are involved in several endangered
species program activities as a result of na-
tional direction from the chief's office.
We have agreed with the Fish and Wild-
life Service (as have all federal land manag-
ing agencies) to a time frame for making rec-
ommendations for the designation of critical
habitat for those species already listed and
for which no critical habitat was established.
This job is in response to the president's
request in his environmental message of 1977
that federal agencies speed up identification
of critical habitats on public lands.
Our regions have been directed to assure
that threatened, endangered, and sensitive
species are adequately covered in regional
and forest land management plans required
by the National Forest Management Act.
Guidelines are being developed to determine
which management direction should be ex-
pressed in regional plans for wide ranging
species and which direction should be left up
to each individual national forest.
The Forest Service will prepare action
plans to accomplish activities identified in re-
covery plans for our agency. Of course, a re-
covery plan cannot commit another federal
agency to the expenditure of funds. Also,
many recovery plans do not provide suf-
ficient details for on-the-ground management
activities, so we must go a step further and
prepare action plans to further refine those
jobs most logical for the Forest Service to ac-
complish, and to serve as our agreement with
the Fish and Wildlife Service to perform cer-
tain tasks in the accomplishment of the re-
covery plan.
We are monitoring, in cooperation with
the states, all populations of threatened and
endangered species on the national forest.
Another program thrust, which is a legal
obligation I have already mentioned, is to re-
view all of our programs and activities and
decide whether or not they may affect a list-
ed species. If the project or activities may af-
fect the species, we formally consult with the
Fish and Wildlife Service.
The final item related to national direction
is to survey listed or sensitive species to lo-
cate populations and define habitat charac-
teristics and biological needs.
Before I talk about some specific projects
for endangered species, I would like to men-
tion our budget and personnel. As you are
aware, the Department of Agriculture gets
no appropriations through the Endangered
Species Act as does the Department of the
Interior and the Department of Commerce.
We do have a specific budget item for en-
dangered species that we make up out of our
normal wildlife appropriations, and then we
have an agreement with Congress about how
much money will be spent on the endangered
species program. This current fiscal year we
are budgeting on the national forest system
$5,223 million for endangered species pro-
grams. I think that this budget is probably
second in size to that of the Fish and Wildlife
Service. I am not sure how large the BLM
budget is. This sounds like a lot of money,
but when you take that much money, allo-
cate it to nine regions, 154 national forest,
and umpteen ranger districts, it is not nearly
as much as it .sounds. In fact, it is not nearly
enough to accomplish a proper job.
The Endangered Species Act, along with
some other legislation, has really changed our
personnel picture also. The Forest Service
during the last four or five years has hired an
162
Great Basin Naturalist Memoirs
No. 3
average of 15 to 20 wildlife biologists a year.
This past fiscal year, we hired 123 biologists
and much of this increased hiring was a di-
rect result of the Endangered Species Act. I
think that upped our total number of wildlife
biologists to somewhere in the vicinity of 370
biologists in the national forest system.
The Forest Service is involved in hundreds
of projects around the country, but these ex-
amples will give you some idea of the type of
things that we are getting into, and some of
the complexities of the situations that we are
dealing with now.
When the California Region began a proj-
ect to identify and recommend critical habi-
tat for bald eagles, they found that not
enough information was available to accom-
plish the job. We knew the habitat conditions
where eagles presently occurred, but infor-
mation was lacking on the criteria for suit-
able imoccupied habitat.
We wanted to designate not only the pres-
ently occupied habitat, but also unoccupied
habitat which was suitable or may be suitable
in the foreseeable future. A program was
started in northern California to gather the
necessary information. A team consisting of a
forester and a wildlife biologist evaluated
every bald eagle nesting territory in the state,
collecting information on such paramaters as
size of tree, aspect, distance from water, dis-
turbance factors, productivity of the nest,
form of the nest tree, timber types immedi-
ately under the nest tree, and timber types
out a certain distance from the nest tree. A
computer program then analyzed the impor-
tant factors that went into making up the
eagle habitat. This program is just being
completed and we are now using the results
of the survey to write criteria for the identi-
fication of bald eagle habitat. Another proj-
ect we are doing with eagles is experimental-
ly improving eagle nest trees. Some trees
have been pruned to improve them for nest-
ing eagles. We have actually tried to encour-
age some eagles to move by judicious prun-
ing of trees and, in some cases, by
constructing artificial nest platforms in the
trees. This is only being done in those areas
where the nest tree is dying or is in an area
that is subject to a large degree of disturb-
ance.
Another project recently completed in
California was the restoration of a peregrine
falcon eyrie. An active nest site on the Men-
dicino National Forest sluffed off of the cliff
face. Climbers went up to the original nest
ledge and made a pattern. The pattern was
then used to preconstruct an artificial nest
platform. Crews then drove metal rods into
the cliff face, installed the artificial nest plat-
form, and covered it with cement and natural
materials to make it look essentially like the
natural nest ledge. As far as we know this has
not been done before, and we are anxiously
waiting to see if the new ledge will be ac-
cepted by the peregrines.
Some interesting work on genetic analysis
with some of the threatened trout and sala-
manders is being done. The Little Kern gold-
en trout occurs only in the Little Kern River
drainage primarily on the Sequoia National
Forest. Over the years, populations of this
threatened species have interbred with in-
troduced rainbow stock so that there are now
very few pure strain Little Kern golden trout
left.
Through the use of the electrophoresis
technique, done under contract with the Uni-
versity of California at Davis, it was deter-
mined exactly which streams within the wa-
tershed contained the pure strain and which
streams were genetically polluted, so to
speak. With this information, agreement was
reached between the California Department
of Fish and Game, the National Park Service,
and the Forest Service on a management
plan for the watershed. This management
plan calls for replacement of many of the
genetically inferior populations with pure
stock, installation of artificial barriers to pre-
vent further interbreeding, and other stream
improvement practices.
The electrophoresis technique was also
used on the shasta salamander, a species list-
ed as rare by the state of California. This
work showed that there were five distinct
populations of this salamander, some of
which had been genetically isolated for well
over 4000 years; these were genetically more
different than some of the full species of sala-
manders were from each other. This brings
up new questions of taxonomy and how spe-
cies should be classified as threatened or en-
dangered and legally protected.
I am going to leave off some of these other
1979
The Endangered Species: A Symposium
163
project examples so that we will have more
time for questions. The Forest Service re-
search arm is completely separate from the
national forest system. It conducts research
on any forest and range land, independent of
ownership. We have 10 work units or work
locations where endangered species work is
going on. This covers about 38 different fed-
erally listed species.
The state and private forestry program is
one which some of vou may not know about.
This third arm of the Forest Service is in-
volved in providing technical advice on re-
source management to state foresters and pri-
vate land owners and administering several
federally financed forestry programs. Of
course, this program is also subject to the En-
dangered Species Act. It is very difficult to
determine the impact of the act on programs
of this type. Both actual and financial assist-
ance and technical assistance given through
the state and private forestry program are
subject to the act.
The National Forest Management Act is
going to drastically change the planning pro-
cesses of the Forest Service. Very briefly,
some of the things that are going to be re-
quired by law now are these: we will set
wildlife goals and objectives, inventory all
species by habitat types, monitor populations
and habitat quantity and quality, quantify
species and habitat diversity, prescribe pro-
tection and management of critical habitats,
and formulate and evaluate alternate man-
agement regimes. These are things that must
be done now by law, and, of course, endan-
gered species management as well as all wild-
life management is tied up in these require-
ments. I will finish with the thought that as
we start making forest plans under the new
National Forest Management Act, we will
most certainly be calling upon you for help.
PANEL: PART III,
THE BUREAU OF LAND MANAGEMENT'S ENDANGERED SPECIES PROGRAM
Richard Vernimen'
Abstract.- It is the responsibihty of the Bureau of Land Management (BLM) to conserve plants and animals . . .
and the habitat on which they depend . . . which are officially listed according to federal or state laws in catregories
that imply significant potential for extinction. The BLM also provides for the conservation of the habitats of unlisted
extinction-prone (i.e., sensitive) plants and animals. It also applies to all BLM programs and actions related to the
public lands, the federal subsurface mineral estate, and the submerged lands of the Outer Continental Shelf (OSC).
The BLM administers 448 million acres of
land within the 11 western states and Alaska
(U.S. Department of the Interior, BLM 1977).
In addition, we are responsible for BLM— au-
thorized actions taking place on the Outer
Continental Shelf and federally owned sub-
surface minerals, i.e., coal, oil and gas, etc.
(hereinafter all of the above lands will be re-
ferred to as BLM-admini,stered lands).
Within these vast acreages and areas of re-
sponsibility we must taken into consideration
the welfare of 48 threatened and endangered
(T/E) animals (U.S. Department of the Interi-
or, BLM 1977) and 3 endangered plants (Fed-
eral Reg. 6/20/78). The T/E plants and ani-
mals occurring on the subsurface and Outer
Continental Shelf (OSC) must also be consid-
ered if BLM-initiated actions affect a T/E
species or its habitat (i.e., oil and gas impacts
on marine mammals). A third category of
species we must take into account are state
T/E species. Our 1977 statistical report listed
138 species of animals.
With the recent passage of the 1978
amendments to the Endangered Species Act
of 1973 (ESA), proposed species must also be
considered for formal consultation. A number
of plants and animals fall into this category.
'Endangered Species Liaison Officer, U.S. Dept. of the Interior, Bureau of Land Management, Washington, DC.
164
Great Basin Naturalist Memoirs
No. 3
Land Use
All actions that we allow on BLM-adminis-
tered lands must be considered for impacts
on threatened and endangered species (T/E),
i.e., oil and gas leases, land exchanges, graz-
ing permits, pipelines, etc.).
The following figures were used for our fis-
cal year 1979 and 1980 budget that shows ac-
tions requiring Section 7 consultation as per
ESA of 1973:
Energy 2500 leases (oil /gas,
coal, geothermal)
Timber Sale of 1.25 billion
board feet
Grazing Issuance of 24,000
use authorizations
Wilderness Completion of 55
studies
State selections 502,900 acres
(excluding Alaska)
Rights-of-way 1700 applications
Mineral leasing (other) 63 million acres
private
290 million acres
other federal lands
Other land actions 8,000 cases
The above are cases or actions readily
identifiable. Each day we encounter new ac-
tions that require review.
Legislation and Authority
Authority-Sources
A. Endangered Species Act of 1973 (16
use 1531 et seq.) as amended.
B. Sikes Act, Title II (16 USC 670 et seq.).
C. National Environmental Policy Act (42
USC 4321 et seq.) as amended.
D. Tlie Federal Land Policy and Manage-
ment Act of October 21, 1976 (P.L. 94-
579).
E. Department Manual 231.1. LA., Gener-
al Program Delegation Director, Bu-
reau of Land Management.
The above acts are our basis for developing
and carrying out an endangered species pro-
gram. The major thrust of our program is
Section 7 compliance and inventory of habi-
tat.
BLM Program
Coordination and Liaison, Section 7 Com-
pliance
Section 7 of the ESA of 1973 directs all
federal agencies on how to comply with the
act. Procedures for this cooperation and con-
sultation can be found in 50 CFR 402 or in
the Federal Register, volume 43, pages
869-876, 4 January 1978.
The major contact on consultation for
BLM is the Fish and Wildlife Service (FWS),
but with our administrative responsibilities
on the Outer Continental Shelf (OCS) we also
consult with the National Marine Fisheries
Service of the Department of Commerce.
Since many of these OCS cases involve the
high seas or foreign countries, we must also
contact the State Department. As you can
see, the Section 7 process can become ex-
ceedingly involved and time consuming.
Because of the mandate placed upon us by
Section 7 of the ESA of 1973, major emphasis
in work load has been shifted to meet it.
Budget increases were added to meet the
need. This is a start, but we are working un-
der pressure to meet the demand because of
other priorities placed upon us, such as the
nation's energy needs.
Critical Habitat Inventory
Tlie president's environmental message of
May 1978 requires that the identification and
determination of "critical habitats" for en-
dangered species be accelerated.
The secretary of the interior is directing
agencies to complete inventories and analyses
for the determinations of critical habitats for
species on their lands by 1 January 1980.
We have 32 of the known species of ani-
mals officially listed on public lands. We
have been given increased funds to complete
this job. Inventories for some species are fair-
ly simple because their respective habiats are
small and centralized. The work begins when
we look at species such as the Bald Eagle or
the American Peregrine Falcon. Habitats of
these species are broad and expansive, requir-
ing many man-hours to complete inventories.
Our participation on recovery teams has
helped to cut this work load down.
1979
The Endangered Species: A Symposium
165
Present Capabilities to Comply with the ESA
of 1973
As of 11 November 1978, the BLM has 249
fisheries and wildlife biologists on board. The
breakdown by numbers and areas is as fol-
lows:
Washington, D.C.
Denver Service Center
Alaska
Arizona
California
Colorado
Eastern States
Idaho
Montana
Nevada
New Mexico
Oregon
Utah
Wyoming
Outer Continental Shelf
Total
6
5
9
18
23
22
3
22
26
22
15
35
22
18
3
249
Within the total 249 biologists, only 2
could be listed as working totally on endan-
gered species, and that is stretching it. We all
have other duties as assigned. I myself func-
tion as the lead in Washington on nongame
species as well as the endangered species liai-
son officer. Mr. Ken Walker, endangered
plant coordinator, will cover the number of
botanists we have working on plants.
Summary
Intensified public concern for our environ-
ment and the flora and fauna within it has
created a demand for all levels of govern-
ment to engage in active and positive pro-
grams to stem the tide of wildlife extinction.
We have embarked on an ambitious program
to protect and benefit endangered plants and
wildlife. Many of our avenues to success are
clouded by complex, competitive demands
on endangered species habitat by other re-
source uses and the nation's need for energy.
Unraveling ecological complexities to isolate
and solve habitat-related problems is not a
simple task. Funding and manpower are not
available to meet all needs. Despite these dif-
ficulties and constraints, we are devoting our
best efforts trying to insure that no additional
plant or animal become either endangered or
extinct on public lands.
Literature Cited
U.S.
U.S.
Department of Commerce, National Oceanic
AND Atmospheric Administration and U.S.
Department of the Interior, Fish and
Wildlife Service. 1978. Interagency coopera-
tion regulations, Endangered Species Act of 1973.
Federal Register 43(2):869-876. January 4.
Department of the Interior, Bureau of Land
Management. 1977. Annual statistical wildlife
report. Unpublished report. Washington, D.C.
1977. BLM statistics for 1976. Washington, D.C.
U.S. Department of the Interior, Fish and Wildlife
Service. 1978. Determination of five plants as en-
dangered species. Federal Register 43:
44810-44812.
PANEL: PART IV,
SUMMARY OF THE ENDANGERED PLANT PROGRAM
IN THE BUREAU OF LAND MANAGEMENT
Kenneth G. Walker'
I'll explain very briefly our function in the
Washington office. You may wonder why
there are two of us here from the Bureau of
Land Management. The primary reason is,
because of the organizational structure at the
Washington office, the responsibility for en-
dangered species coordination is in the Divi-
sion of Wildlife, with Dick Vernimen as the
coordinator for the Bureau of Land Manage-
ment. My function in the Division of Water-
shed is to assist or carry on the coordinating
role for endangered plant species. The sym-
'Endangered Plant Coordinator, U.S. Dept. of the Interior, Bureau of Land Management. Washington, D.C.
166
Great Basin Naturalist Memoirs
No. 3
posium, I feel, has been very enlightening.
The scientific community in many instances
seems to be at odds as to what really needs to
be done for endangered species, what the
needs are, and what the protection systems
should be. We in the federal agencies do not
have many options, although we have our
opinions. Our options are limited to the
methods for which we follow the dictates of
legislation.
Policy for endangered plant species is very
similar to that described by the Forest Ser-
vice. Our prime effort is not only to protect
and conserve listed species, but also to carry
it a step further and to protect and conserve
the proposed species with the idea that if we
can manage these species and their habitat
the situation will be avoided where they will
require official listing. We recently devel-
oped a policy for endangered species which I
will summarize. It is the policy to protect,
conserve, and manage federally and state-list-
ed or proposed listings of sensitive, endan-
gered, or threatened plants and to use its au-
thorities in furtherance of the purposes of the
Endangered Species Act and similar state
laws. The bureau, through its actions in all
planning and management activities, will in-
sure that the actions authorized, funded, or
carried out will not jeopardize the continued
existence of such species or result in the de-
struction or modification of the critical habi-
tats. To summarize the policy, as the Forest
Service mentioned, our intent is to not only
follow the letter of the law, but also the spirit
of the law. We have issued several guidelines
to our field office to follow this policy. In
doing this, we have asked our field office to
do two things: first, to add each candidate or
listed species which is known or expected to
occur within their area of responsibility to a
list of these species that will be developed
and maintained by our state directors within
the area of jurisdiction. The area of responsi-
bility in Utah, for example, would be the en-
tire state, which in tvirn requires a lot of
coordination with the universities, state
agencies, private concerns, and others, wher-
ever we can acquire the interest. A second
appeal would be for state directors to deter-
mine those species which are known or sus-
pected to occur on bureau-administered lands
or can reasonably be expected to be in-
fluenced by bureau actions. The Bureau of
Land Management has the responsibility for
management of surface areas, but there also
are many areas where we have responsibility
for the subsurface minerals management.
Coal, in Utah, is an example where we man-
age the subsurface minerals but, we do not
own the surface. This creates many problems.
I will now summarize the program status
for the endangered species program in the
BLM. I feel almost embarrassed sitting by the
Forest Service people when they talk about
their funding levels. Our funding for endan-
gered plant species has not been a direct
fimding effort. We've acquired from other
programs approximately $400,000. This in-
cludes partial funding of about 40 personnel.
Unfortunately, not very many of them are
able to spend their full-time in the endan-
gered species effort. We do have a few full-
time botanists. The endangered plant pro-
gram in this bureau is viewed as low priority
because of its magnitude. On public lands,
only three species have officially been listed.
All three of them are in California. We have
several hundred proposed species located on
public lands. Our endangered plant species
program is primarily, at least at this time, as
Duane Atwood mentioned this morning, in
the inventory stage. We're not yet to the
point where we're really able to prepare or
do active planning for a particular species or
a particular group of species. Our efforts are
tied rather closely to our Environmental
Statement (ES) Program in the bureau, par-
ticularly the range program, which is a mag-
nanimous effort. We have several hundred
environmental impact statements to prepare
within the next few years. Our endangered
species inventory efforts have pretty much
centered around ES efforts. Our efforts and
methods in conducting these inventories are
varied. Some are done in-house by our own
people. Many of them we are able to conduct
through contracts with universities and oth-
ers who have such capability.
1979
The Endangered Species: A Symposium
167
Questions to the Panel
Q. The Endangered Species Act is rather narrow about
defining this problem. There are quite a few other
programs that can be appHed. Many of the federal
land agencies have natural area programs. There are
also a number of wildlife programs that can be
brought to bear on the question of peripheral species
and their distribution. We have the same problem
with plant distribution, so I'm not sure the endan-
gered species program is the right place for that
kind of program, depending, of course, on what hap-
pens to the whole range. There are a lot of other
programs that could help there.
A. That particular problem is one of the things we are
trying to address with our sensitive species category
in our total endangered species program. We can
take species like this and put them on our sensitive
species list and then apply land management prac-
tices or management practices in a special way.
There won't be the legal requirements, but we
would treat them for land management purposes the
same way we would treat a legally listed species.
(Mcllwain)
Q. I have a correlary to this I need to address. I don't
think it's been addres.sed to the extent that I need to
understand it. Having worked for a private con-
sulting firm, I've often been caught between two
grist mills of state species lists and also federal spe-
cies lists. Specifically, I'd like to know what your
plans are for the future. I don't think I understand
how you're going to correlate and work out these is-
sues with the states. For example, the Hamper Proj-
ect is not ad ministered by the state. It's a national
environment research park. What if we have a spe-
cies there that is peripheral and we want to protect
it, but the State of Washington doesn't. The popu-
lation is found in Washington and parts of Oregon,
Idaho, and Utah, but in most areas it doesn't war-
rant or merit consideration as a threatened species.
How are you going to handle this conflict with the
states? Will you be able to support it?
A. Well, as a matter of fact, I don't see any conflict
with the states at all in a situation like this. If a giv-
en state has its own endangered species legislation,
and if a particular species, be it a peripheral species
or whatever, is in trouble in that state, I see nothing
wrong with that state listing that species under its
legislation as an endangered species and protecting
it accordingly. (Mcllwain)
Q. By a conflict, I mean to be able to fimd them and
support them financially. Most of the states don't
have an adequate threatened and endangered spe-
cies program, especially from the standpoint of fimd-
ing resources. You have infinite amounts compared
to what most of them do. Will you be able to sup-
port them on the basis of those peripheral popu-
lations?
A. We have two separate fimding resources in the en-
dangered species program. One is the Section 1.5
monies, which our general appropriation authorizes,
and the other is the Section 6 money, which is dedi-
cated specifically to a grant and aid program
through cooperative programs with the states. We
have not, as a matter of fact, been able to obligate
that money as quickly as we would like to— simply
because there has not been enough demand in the
states to really get with the program. I don't see any
difficulty in fimding through a matching 66 percent
federal share-.3.3 percent state share for state activi-
ties. I don't think we're going to run out of money
any time sooner. (Spinks)
Q. Wouldn't those matching fimds work only for spe-
cies that are listed as endangered species under the
federal act?
\. No, if they're considering it for listing under the
state act. They would also be eligible for funding.
(Spinks)
Q. I've enjoyed very much your program, but you have
not mentioned the aquatic forms. Now you take the
fisheries on the North Atlantic, the whaling. They're
vital problems with which we must deal. It seems to
me that not only will we have to be financed, but it
may even be we'll have to use a little military
strength to restrain some of these people who say
they have a right to hunt a particular species, the
whale and so on. That is a major problem as I see it
in connection with the immediate approach in deal-
ing with these species.
.\. Your point is well taken. I'm glad the National Ma-
rine Fisheries Service is in this act with us. There is
basically a division of responsibility in the act be-
tween the Departments of Commerce and the Inte-
rior, and the oceanic species are under the pro-
tection and administrative authority of the National
Marine Fisheries Service. Certainly we do not in any
way want to diminish the value of those species, as
vou point out, but that is again the prerogative of
the National Marine Fisheries Service; and, as Mr.
Vernimen mentioned, the Bureau of Land Manage-
ment under the OCS leasing program does become
involved with the National Marine Fisheries Service
in the consultation process, like considering such
species as the bow-head whale in Alaska, for in-
stance. (Spinks)
Q. I have a comment on a previous question. The State
of Washington is being fimded now by endangered
species dollars to come up with a list of the state's
threatened and endangered species, so it is possible
to do that. The state game department is involved in
that.
Q. My question to you managers is from the point of
view of private industry. I'm a representative of
Utah Power and Light Company, and I'm not a biol-
ogist. I've learned a lot here in the last couple of
days about biolog\'. Obviously, the vital question to
us is this. We realize that recent amendments to the
act have created a lot of work for you guys to do.
.\re we going to have to wait for you to get all this
work done before we can build any new plants, or
will we have to provide some of the fimding to get
some work done on a specific basis by ourselves?
.\. No, you do not, as a matter of fact, have to wait un-
til there are new Section 7 regulations promulgated,
which could take some time. We are proceeding
with the consultation process under the existing Sec-
tion 7 regulations which Jerry Mcllwain alluded to
as having been published in January 1978. The
168
Great Basin Naturalist Memoirs
No. 3
world is not going to stop until we have the new
regulations. (Spinks)
Q. I want to ask a question concerning the program of
the Forest Service people and the BLM in terms of
the protective habitat, just to clarify what I'm con-
cerned with. For a number of years I cooperated
with some of the folks from California who were
trying to preserve some sand dunes in southeastern
California, southern Nevada, and perhaps other
areas from dune buggies and off-road vehicles that
just traversed the area without any concern for the
animals or the plants that were there. Now I haven't
heard from Bob Stebbins or Dave Wake or some of
those folks for a few years as to whether or not they
have succeeded in convincing the Bureau of Land
Management that some steps should be taken to pro-
tect those sand dunes habitats before the sand adapt-
ed and a number of other forms are exclusively re-
stricted to those areas. What has been done and
what is the program of the Bureau now to protect
habitat from these kinds of degradations?
A. We do now have the three species in California that
are officially listed. I believe two of them are in the
sand dunes area. For one of them, specifically, the
Eureka Sand Dune Grass, the Bureau of Land Man-
agement has tried to close this area. We've received
some criticism as to how effective the closure of
these lands has been. Others say it's been very effec-
tive. But, to go back to the other part of my answer,
our planning process is to go through our inventories
and identify critical habitats, sensitive species, pro-
posed species with their habitats, and, through the
planning process, tie these areas in with other pro-
posed actions, one of which could be off-road vehicle
use. Then, in the final recommendations through our
planning process, the decision is made then as to
what action will be taken in regards to that area—
whether it be closure, restrictions from other uses,
grazing, off-road vehicles, or other means to protect
certain species. This is the process. Now the actual
implications of success to this process we've yet to
see in many cases, but we are making a sincere at-
tempt. (Walker)
1 think the other area we can talk about, speaking
of California, is the Desert Tortoise area, on which
Dr. Kristine Berry and a team of other people have
been working. We have fenced out most of that
area. We have also posted signs, although I have
heard recently that 400 signs have disappeared. We
also have off-road vehicle regulations we are looking
at, where we would close it to such vehicles. In-
cidentally, one of the beetles proposed does occur
right in the middle of an off-road vehicle area in Ne-
vada.
We also have authority for emergency closures if
we want to use it. (Vemimen)
Q. What I'm trying to suggest is that if the Bureau of
Land Management or private industries, do not pro-
tect the desert habitat, we stand to lose a lot of this
very valuable material.
Q. I'd like to bring up the controversy of reintroduction
in an area of historic range, but not now pre.sent.
We ran into it with the Colorado squawfish. I was
wondering if the land management people would
comment about taking an endangered species into a
recovery plan, trying to get it off the list more or less
by reintroduction into the historic range. Do you nm
into the resistance of a local forester or a local dis-
trict manager saying, "If I have to worry about that
I won't be able to go into the campground"?
A. That's a very difficult and subjective question, one
which is extremely hard to formulate a policy on be-
cause you have to adjust to the situation on some-
thing like that. Certainly we're not going to reintro-
duce grizzlies to the plains where they once
occurred around the Denver area. That's completely
unreasonable. On the other hand, in the process of
identifying the essential habitats or the legally desig-
nated critical habitats on the public lands, we found
a lot of these that are historical into which we can
logically expand species. Somewhere in the middle
between the unreasonable and the feasible is the
line, and how you define that line is very difficult.
It's going to be a subjective decision. (Mcllwain)
I'd like to cite an example. In Arizona they want
to reintroduce the woundfin into historical habitat.
At the same time, this habitat is the number one
geothermal exploration area in Arizona. This is the
type of administrative problem we get into, and I
am to the point now where I tend to agree with a
state director who says, "No, not until further stud-
ies are completed." The problem is "Can we under
the act say no? " So, right now that opinion is in the
solicitor's office. These are the kinds of things you
nm into. You've got to use some judgement. We
have an area that's being managed for some specific
resource and then all of a sudden we throw some-
thing else in there that is going to change it. We're
going to have to weigh that very heavily before we
reintroduce it. (Vernimen)
I'd like to make one more comment before we
beat this question to death. Is this a situation where
it is really necessary for the survival of the species,
or is it something we would like to see for the pro-
mulgation of the species? To me this is the big ques-
tion, and it gets down to whether we really need to
or just want to. I think reintroduction of a species
should be considered as a last resort in the perpetu-
ation of the species. We have to consider the prob-
lems we nm into with reintroduction. Are we creat-
ing more problems than we are solving?
Q. In Utah we have watched the systematic destruction
of the Lynndyl Sand Dune area, the Coral Pink Sand
Dime area, and the Hurricane Sand Dime area, all of
these under major control of the Bureau of Land
Management. I am about to describe a new species
of sunflower from the Lynndyl Sand Dunes, known
in Utah by the misnomer. Little Sahara. It is not. It
cannot be. It is systematically being destroyed.
We're not talking about reintroducing something,
but we're talking about protecting something the
Lynndyl Sand Dunes have, among other unique spe-
cies which Professor Stutz mentioned earlier today.
The Coral Pink Dimes have still others. The ones at
Hurricane are unexplored. We don't know what's on
them. We may never be able to find out because of
off-road vehicle use. What is the potential then, for
a turnabout for at least a part of these areas?
1979
The Endangered Species: A Symposium
169
A. It just so happens that when I was in the Richfield
district, as well as being a wildlife biologist I was a
recreation specialist and I did have something to do
with Little Sahara as you call it. I am not too famil-
iar with the Hurricane area you talk about. Now the
southern part of the Coral Pink Sand Dunes— correct
me if I'm wrong— are managed by the state as a state
park. My question is "Have you contacted the state
office here and informed the Bureau of Land Man-
agement that you have found those plants?" (Verni-
men)
Q. How does the BLM treat endangered or threatened
species on subsurface land? By that we mean private
ownership of the surface and someone else owns the
minerals, oil, gas, coal, etc.
A. First of all, the identification of the critical habitat
and the inventories (unless we have an action taking
place right at that time) is the responsibility of the
Fish and Wildlife Service on the private lands. If
you take the case of the Red Cockaded Woodpecker
in Alabama, where the BLM has some subsurface
coal, the BLM is doing the inventories. The BLM is
also doing the inventories on the Eastern Cougar.
We are in the process of contracting an individual to
do the inventories on that. If we would let a lease
go, we are initiating an action. We are responsible
to see that that species is protected.
Q. Is that also the case for critical habitat on state land
for endangered plants and endangered plants on pri-
vate lands?
A. Are you saying designation of a critical habitat or
protection of a critical habitat? (Vernimen)
Q. Identifying of an endangered plant on private sur-
face land but federal subsurface. Wouldn't the pri-
vate landowner have the discretion of saving that
plant?
A. Well, no. If we didn't sell the coal in there, it
wouldn't be mined. (Vernimen)
Q. So you could deny the lease of such materials?
A. That's correct. (Vernimen)
A. May I address a couple of things that you said. Num-
ber one, plants are not protected from being taken
under the act. If the private landowner has a bunch
of furbish louseworts or whatever and the man
wants to go out there and chop them all down with
a hoe, that's legal. The second point is that, in terms
of having something protected by virtue of having
critical habitat determined for it, it is protected
from a federal action under Section 7, whether or
not there is any critical habitat there. There is a bas-
ic question of jeopardy, .\mong other things in Sec-
tion 7, besides almost an affirmative action clause
for federal agencies to do some good things for listed
species, there is the no section that says they shall
insure their actions do not jeopardize the continued
existence of a species. So, with or without critical
habitat designation, there would still be this respon-
sibility to not jeopardize the species. (Spinks)
Q. You said yesterday, when you were ennumerating
the amendments to the act, that the application for
critical habitat would be withdrawn.
A. Our understanding at this point in time is that the
outstanding proposals for critical habitat designation
will be withdrawn and reproposed to bring them in
compliance with the 1978 amendments. (Spinks)
Q. In response to that, I'd like to ask Mr. Mcllwain
what kind of protection will be given to the critical
habitats of the grizzly bear, mainly because there is
such a controversy over how much should be given
them?
A. As far as I'm concerned, critical habitat on forest
service lands doesn't really mean very much because
we're protecting that critter or the habitat of that
listed species as a requirement of the law regardless
of whether it is legally designated as critical habitat
or not. We have management programs established
now to protect grizzly bear habitat and we're estab-
lishing others as time goes on. It really makes little
difference whether critical habitat is legally desig-
nated or not for the time being. (Mcllwain)
Q. I'd like a little clarification with regard to the con-
flict between the Endangered Species Act and min-
ing development. Tliere seems to be a rather ne-
bulous area.
A. I know just what you're talking about. I have several
memos in my office about people asking just where
does the 1872 Mining Law and the Endangered Spe-
cies Act fit in. As you know, they are both non-
discretionary, and it's kind of like two penalties on a
football field. They more or less nullify one another.
I'm not at liberty to comment right now. The solic-
itor is coming out with an opinion on the 1872 Min-
ing Law and the Endangered Species Act, and I
don't know yet what he is going to say. Right now
they can go ahead with exploration and mining de-
velopment for hardrock minerals, gold, silver, and so
forth. There is nothing that the Endangered Species
Act can do to .stop them. Nothing. (Vernimem)
My only comment is that we may be finding out
what happens in this regard before too long because
we have two situations now on Forest Service land,
two similar conflicts, one in Arizona and one in Cali-
fornia, conflicts between the Mining Act of 1872 and
the Endangered Species .^ct in relation to an appli-
cation for mining within a bald eagle nesting terri-
tory. Either or both of those may get to court before
too long. (Mcllwain)
Q. Would the Forest Service get a different opinion if
you went through a different group as it were?
A. Well, we go through a different solicitor. We go
through the USDA Office of the General Council,
which is the same as a solicitor. (Mcllwain)
Q. Are you seeking an opinion also?
A. No, we're not. (Mcllwain)
The bottom line here on the opinion of a solicitor
or the Office of General Council from the Depart-
ment of Agriculture, as in the case of the U.S. Forest
Service, is an internal guidance mechanism for that
department or agency. The real bottom line is writ-
ten through the development of case law, and, until
there is sufficient litigation involving such conflicts
as mining and the Endangered Species .Act, there
will not be a hard and fast answer to that very good
question. (Spinks)
Q. Your statement puzzles me a little bit regarding con-
flict between the Endangered Species Act and the
mining law with respect to bald eagles, especially
the protection of bald eagles is .so stringent with re-
gards to nesting areas, etc. Isn't the Forest Service
required to adhere to that?
170
Great Basin Naturalist Memoirs
No. 3
A. Yes, we're required to adhere to that, but there is a
question as to when you are really harrassing a bird.
In the particular conflicts that I'm talking about, we
have established a territory for a bald eagle nesting
pair, and the mining people want to build a road
through that territory and mine outside of it. We've
told them no. We're set up to be sued any way we
go. If we give a permit to the mining operation,
we're going to be sued by the environmentalists un-
der the bald eagle act or the Endangered Species
Act or others. On the other hand, if we say no, we'll
be sued by the mining interests. In this particular
case we decided to remain on the side of the envi-
ronmentalists. (Mcllwain)
Q. The Fish and Wildlife Service just recently issued a
proposal for critical habitat for the squawfish. Will
you finalize that rule making, or are you still work-
ing that thing over? What is that status.
A. Like other proposed rule makings for critical habitat
determination, that will have to be reproposed to
comply with the 1978 act amendments.
Q. It will be reproposed then at some future date?
A. Yes it will. (Spinks)
AUTHOR AND TITLE INDEX FOR THE ENDANGERED SPECIES: A SYMPOSIUM
Atwood, Duane, article by, p. 81.
Baumann, Richard W., article by, p. 65.
Clement, Roland C, article by, p. II.
Culture and species endangerment, p. II.
Day, Douglas, article by, p. 35.
Deacon, James E., article by, p. 41.
Endangered and threatened fishes of the
West, p. 41.
Endangered and threatened plants of Utah: A
case study, p. 69.
Endangered animals in Utah and adjacent
areas, p. 35.
Endangered species: Costs and benefits, p.
151.
Endangered species on federal lands. Part I:
Introduction, p. 159.
Endangered species on federal lands. Part II:
Forest Service philosophy of endangered
species management, p. 159.
Endangered species on federal lands. Part III:
The Bureau of Land Management's endan-
gered species program, p. 163.
Endangered species on federal lands. Part IV:
Summary of the endangered plant pro-
gram in the Bureau of Land Management,
p. 165.
Harper, K. T., article by, p. 129.
Holmgren, Arthur H., article by, p. 95.
Introductory remarks, p. 1.
Lovejoy, Thomas E., article by, p. 5.
Management programs for plants on federal
lands, p. 81.
Mcllwain, Jerry P., article by, p. 159.
Murphy, Joseph R., article by, p. 1.
Perspective, p. 17.
Pister, Edwin P., article by, p. 151.
Rare aquatic insects, or how valuable are
bugs? p. 65.
Rare species as examples of plant evolution,
p. 113.
Some reproductive and life history character-
istics of rare plants and implications of
management, p. 129.
Spencer, Donald A., article by, p. 25.
Spinks, John L., articles by, pp. 17, 159.
Stebbins, G. Ledyard, articles by, pp. 87, 113.
Strategies for the preservation of rare ani-
mals, p. 101.
Strategies for preservation of rare plants, p.
95.
Strategies for preservation of rare plants and
animals, p. 87.
Stutz, Howard C, article by, p. 119.
Tepedino, V. J., article by, p. 139.
The epoch of biotic impoverishment, p. 5.
The importance of bees and other insect pol-
linators in maintaining floral species com-
position, p. 139.
The law and its economic impact, p. 25.
The meaning of "rare" and "endangered" in
the evolution of western shrubs, p. 119.
Vernimen, Richard, article by, p. 163.
Walker, Kenneth G., article by, p. 165.
Welsh, Stanley L., article by, p. 69.
White, Clayton M., article by, p. lOI.
171
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