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Full text of "Great Basin naturalist memoirs"

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. 

The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been published 
from one to four times a year since then by Brigham Young University, Provo, Utah. In gener- 
al, only previously unpublished manuscripts of less than 100 printed pages in length and per- 
taining to the biological and natural history of western North America are accepted. The 
Great Basin Naturalist Memoirs was established in 1976 for scholarly works in biological natu- 
ral history longer than can be accommodated in the parent publication. The Memoirs appears 
irregularly and bears no geographical restriction in subject matter. Manuscripts are subject to 
the approval of the editor. 

Subscriptions. The annual subscription to the Great Basin Naturalist is $12 (outside the 
United States $13). The price for single numbers is $4 each. All back numbers are in print and 
are available for sale. All matters pertaining to the purchase of subscriptions and back num- 
bers should be directed to Brigham Young University, Life Science Museum, Provo, Utah 
84602. The Great Basin Naturalist Memoirs may be purchased from the same office at the 
rate indicated on the inside of the back cover of either journal. 

Scholarly Exchanges. Libraries or other organizations interested in obtaining either journal 
through a continuing exchange of scholarly publications should contact the Brigham Young 
University Exchange Librarian, Harold B. Lee Library, Provo, Utah 84602. 

Manuscripts. All manuscripts and other copy for either the Great Basin Naturalist or the 
Great Basin Naturalist Memoirs should be addressed to the editor as instructed on the back 



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 








6 


6.7 


5 


4.4 








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 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 1 


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 


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 





7 





.\sh Spring 


5 





8 





Crystal Spring 










Hiko Spring 


1576 





6 





Mormon Spring 










Preston Town Spring 










Preston Big Spring 














1.58 





11 





11 







106 





64 











20 





64 





83 







54 





52 





58 
17 







1979 



The Endangered Species: A Symposium 



51 



70. 


• 


• above narrows 
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 





920 


5.1 






11 9 


11 
01 










4 
82 






16 25 

25 

17 



68 



15 


1259 


253 





90 


10 224 9 


5 







2828 .04 356 





69 


313 20 


27 


59 


159 


1051 


188 





5 


25 






20 


704 


440 






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 







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. 

Literature Cited 

Clement, D. A. 1979. Rare species and culture. Great 
Basin Nat. Mem. 3:11-16. 

CoTTAM, W. P. 1961. Our renewable wild lands— a chal- 
lenge. Univ. Utah Press, Salt Lake. 

Cross, J. N. 1975. Ecological distribution of the fishes of 
the Virgin River (Utah, Arizona, Nevada). Un- 
published thesis. Univ. Nevada, Las Vegas. 

1978. Status and ecology of the Virgin River 

Roundtail Chub, Gila robusta seminuda (Os- 
teichthyes:Cyprinidae). S.W. Nat. 23(3): 519-528. 

Deacon, J. E., and M. S. Deacon. 1979. Research on 
endangered fi.shes in the national parks with spe- 
cial emphasis on the Devils Hole pupfish. Proc. 
First Conf. on Scientific Research in the National 
Parks. Vol. 1:9-20. 

Deacon, J. E., C. Hubbs, and B. J. Zahuranec. 1964. 
Some effects of introduced fishes on the native 
fish fauna of southern Nevada. Copeia. 
1964(2):384-388. 

Deacon, J. E., G. Kobetich, J. D. Williams, and S. 
Contreras. 1979. Fishes of North America, en- 
dangered, threatened or of special concern: 1979. 
Fi.sheries 4(2):29^4. 

Deac;on, J. E., and W. L. Minckley. 1974. Desert fish- 
es, pp. 385-487. In: Desert Biology, Vol. 2, Aca- 
demic Press, N.Y. 

Hastings, J. R., and R. Turner. 1965. The changing 
mile. Univ. Arizona Press, Tucson. 

Hubbs, C, and J. E. Deacon. 1964. Additional in- 
troductions of tropical fishes into southern Ne- 
vada. S.W. Nat. 9(4):249-251. 



64 



Great Basin Naturalist Memoirs 



No. 3 



HuBBs, C. L., R. R. Miller, and L. Hubbs. 1974. Hydro- 
graphic history and relict fishes of the north-cen- 
tral Great Basin. Mem. California Acad. Sci. 
7:1-259. 

HoLDEN, P. B., AND C. B. Stalnaker. 1975a. Distribu- 
tion of fishes in the Dolores and Yainpa river sys- 
tems of the upper Colorado Basin. S.W. Nat. 
19(4):403-412. 

1975b. Distribution and abundance of main- 
stream fishes of the middle and upper Colorado 
River basins, 1967-1973. Trans. Amer. Fish. Soc. 
104(2):217-231. 

Iltis, H. H. 1967. To the taxonomist and ecologist 
whose fight is the preservation of nature. Bio- 
science 17(12):886-890. 

LocKHART, J. N. 1979. Ecology of the woundfin minnow, 
Plagopterits argentissimiis Cope. Unpublished 
thesis. Univ. Nevada, Las Vegas. 

LovEjOY, T. E. 1979. The epoch of biotic impover- 
ishment. Great Basin Nat. Mem. 3:5-10. 

McNatt, R. 1979. Fish habitat loss in the San Pedro 
River, Arizona. Proc. Desert Fish. Coun. In press. 

Miller, R. R. 1959. Origin and affinities of the fresh- 
water fish fauna of western North America. In: 
C. L. Hubbs, ed., Zoogeography. Amer. Ass. Adv. 
Sci., Washington, D.C., Publ. No. 51, pp. 
187-222. 

1961. Man and the changing fish fauna of the 

American Southwest. Pap. Michigan Acad. Sci., 
Arts and Lett. 46:365-404. 

Miller, R. R., and J. R. Alcorn. 1946. The introduced 
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 









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 





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 



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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 



111 



ities tor the endangered species program. Trans. 
N. Am. Wildl. Natl. Res. Conf. 40:142-156. 

Spellerberg, I. F. 1975. Conservation and management 
of Britain's reptiles based on their ecological and 
behavioral requirements: a progress report. Biol. 
Conserv. 7:289-300. 

Springer, P. F., G. V. Byrd, and D. W. Woolington. 
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threatened species. Univ. of Wisconsin Press, 
Madison. 

Sullivan, A. L., and M. L. Shaffer. 1975. Biogeo- 
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Te.mple, S. a. (ed.). 1978. Endangered birds: manage- 
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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 



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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 

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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